NOAA Technical Report NMFS 103

Benthic Macrofauna of the

New York Bight, 1979-89

Robert N. Reid

David J. Radosh

Ann B. Frame

Steven A. Fromm

U.S. Department of Commerce

December 1991
NOAA Techncial Report NMFS

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NOAA Technical Report NMFS 103

Benthic Macrofauna of the

New York Bight, 1979-89

Robert N. Reid

David J. Radosh

Ann B. Frame

Steven A. Fromm

December 1991

U.S. DEPARTMENT OF COMMERCE

Robert Mosbacher, Secretary

National Oceanic and Atmospheric Administration

John A. Knauss, Under Secretary for Oceans and Atmosphere

National Marine Fisheries Service

William W. Fox Jr., Assistant Administrator for Fisheries
The National Marine Fisheries Service (NMFS) does not approve,

recommend or endorse any proprietary product or proprietary

material mentioned in this publication. No reference shall be made

to NMFS, or to this publication furnished by NMFS, in any advertising or sales promotion which would indicate or imply that NMFS

approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose

an intent to cause directly or indirectly the advertise9 product to

be used or purchased because of this NMFS publication.

-------------------"
Benthic Macrofauna of the New York Bight, 1979-89

ROBERT N. REID, DAVID J. RADOSH

ANNE B. FRAME, AND STEVEN A. FROMM

National Marine Fisheries Service

Northeast Fisheries Science Center

Sandy Hook Laboratory

Highlands, NewJersey 07732

ABSTRACT

The benthic macrofauna of the New York Bight has been monitored extensively, primarily to determine trends over space and time in biological effects of waste inputs. In

the present study, from 44 to 48 stations were sampled each summer from 1980-1985.

Data from other Bight benthic studies are included to· extend the temporal coverage

from 1979 to 1989. Numbers of species and amphipods per sample, taken as relatively

sensitive indicators of environmental stress, showed consistent spatial patterns. Lowest

values were found in the Christiaensen Basin and other inshore areas, and numbers

increased toward the outermost shelf and Hudson Shelf Valley stations. There were

statistically significant decreases in species and amphipods at most stations from 1980 to

1985. (Preliminary data from a more recent study suggest numbers of species increased

again between 1986 and 1989.) Cluster analysis of 1980-85 data indicated several distinct

assemblages-sewage sludge dumpsite, sludge accumulation area, inner Shelf Valley,

outer Shelf Valley, outer shelf-with little change over time. The "enriched" and "highly

altered" assemblages in the Basin appear similar to those reported since sampling began

there in 1968. No consistently defaunated areas have been found in any sampling programs over the past 20 years. On a gross level, therefore, recent faunal responses to any

environmental changes are not evident, but the more sensitive measures used, i.e. numbers of species and amphipods, do indicate widespread recent effects. Causes of the

faunal changes are not obvious; some possibilities, including increasing effects of sewage

sludge or other waste inputs, natural factors, and sampling artifacts, are discussed.

Introduction _

The "apex" (northwest corner) of the New York Bight

(Figs. 1, 2) has long received large organic carbon

and toxicant inputs from the Hudson-Raritan estuary

(since the 1800s) and from dumping of dredged material (since 1914) and sewage sludge (1924), with

smaller inputs from several other sources (Mueller et

al. 1976; New York City Department of Environmental

Protection 1983; Stanford and Young 1988). Fates

and biological effects of the introduced contaminants

have been studied extensively; see Gross (1976) and

Mayer (1982) for symposia reviewing Bight studies.

Among the objectives have been to determine overall

contaminant influences, to partition influences

among the various sources, and to detect changes

over time in fates and effects.

This report concentrates on benthic macrofau.na as

indicators of environmental change in the Bight.

Benthic macrofauna (hereafter called "benthos,"

meaning the bottom-living invertebrates collected in

"grab" samples and retained on sieves of 0.5-1.0 mm

mesh size) are often used in this context. Their relative immobility and intimate association with contaminant-accumulating sediments make them among the

best biological indicators of effects of contaminants

and of environmental change in general (Kuiper

1986; Jackson and Resh 1989). The benthos is also

monitored to assess actual or potentia! importance of

an area as a food and contaminant source for bottomfeeding resource species. Some discussion of the

Bight benthos in these contexts is given elsewhere

(e.g., Boesch 1982; Steimle 1985).

There have been at least 17 major sedimentbenthos sampling efforts in the Bight, beginning in

1966 (see Reid and Steimle [1988] for a summary of

existing surveys). The largest was NOAA's Marine

Ecosystems Analysis (MESA)-New York Bight

1
2 NOAA Technical Report NMFS 103

Project, with sampling from 1973-1976. One of the

products of the MESA program was a recommendation for continued monitoring of several components of the Bight's ecosystem, including sediments

and benthos. The monitoring plan proposed was described in Reid et al. (1982). The sediment-benthos

portion of the plan was modified somewhat (see

Methods), and annual sampling was conducted

from 1980 through 1985.

This report presents data collected on macrobenthic species richness, numbers and biomass of

dominant species, numbers of amphipods, and similarities of community structure over space and time.

Raw data are available on request. The report also

includes relevant data from NOAA's Northeast Monitoring Program (NEMP) (Reid et al. 1987), and initial data from a 1986-89 study of responses of the

benthos to the phaseout ofsewage sludge dumping in

the Bight apex (Environmental Processes Division,

Northeast Fisheries Center 1988, 1991). Data from

this report provide baselines against which responses

to phaseout can be measured. These data may also be

useful in guiding future decisions such as where to

relocate the present dredged material dumpsite and

where and how to mine sand to minimize impacts on

benthos (and potentially on resource species which

interact with benthos).

Methods --------------

The basic monitoring scheme involved sampling 44 to

48 stations (Figs. 1, 2) each summer from 1980

through 1985. The annual samplings were conducted

on 28 July-5 August 1980, 10-19 August 1981, 9-15

September 1982, 19-21 July and 22 August-7 September 1983, 21-31 August 1984, and 30 September5 October 1985 (Table 1).

Six of the stations (4, 6, 7,15,26, and 31) were also

part of the NEMP benthic program. These stations

were sampled semiannually beginning in December

1979, and resulting data are included to extend the

study's temporal coverage. For the same reason, preliminary data from a more recent study of benthic

responses to the phaseout of sewage sludge disposal

in the inner Bight are also presented. Stations 6 and

11 of the annual monitoring surveys were used as the

"sludge accumulation" and "ref~rence" stations, respectively, for the phaseout study. July, August, and

September 1986-1989 samplings at these stations are

also included to extend the report's temporal coverage (only data on numbers of species and of Capitella

spp. for 1988, and preliminary data on species numbers for 1989, are available) (Environmental Processes Division, 1991).

In the annual monitoring survey, one sample per

station was taken in 1980 and 1981, and two samples

in 1982-85. The NEMP program took five samples

per station, and the phaseout study, three samples.

Table 1 gives dates on which individual stations were

sampled, as well as numbers of samples analyzed, station locations and depths, and sediment grain sizes,

carbon and nitrogen contents.

Loran C, with a nominal accuracy of ::1::50 m

(Holme and McIntyre 1984), was used to locatestations. Samples were taken with a 0.I-m2 SmithMcIntyre grab. Plastic tubes of 2.7-cm inner diameter

were used to take one subsample from each grab

for analysis of sediment grain size, organic carbon

and nitrogen, and one subsample for heavy metals.

The remainder of each grab was rinsed through a

0.5-mm sieve. Retained materials were fixed immediately in 10% buffered formalin with Rose Bengal biological stain, and were transferred to 70% ethanol

with 5% glycerin one to three days later. After storage

for at least six months, samples were sorted using dissecting microscopes. Identifications were to species

level whenever possible, except for rhynchocoels.

Oligochaetes, archiannelids, and colonial forms were

not enumerated owing to uncertainty of identification and/or difficulty of quantification. Wet

weight biomasses were determined by blot-drying

each taxon on absorbent toweling for 3 minutes and

weighing to the nearest milligram on an electronic

balance.

Spatial contours of numbers of species and amphipods were drawn using Surface II software (Sampson

1978). In an exploratory attempt to describe trends

in numbers of species and amphipods over time, linear regressions were calculated from the species and

amphipod versus time data. The data are not ideally

suited for linear regression analysis, for several reasons. Changes in the "dependent" variables, numbers

of species and amphipods, are not actually caused by

changes in the "independent" variable, time. There

was no a priori hypothesis except for the implicit one

that numbers of species and amphipods would not

change over the course of the study unless there were

underlying environmental change (the analysis does

not address the processes behind any trends detected). A linear fit to the data may not best represent

the underlying processes, but exploring non-linear

fits may not be justified when those processes are not

postulated a priori. However, the analysis does provide

an objective way of,describing and comparing trends.

Time series or trend analysis could not be used
Table I

Location, depth, mean grain size, carbon and nitrogen, and numbers of benthic samples analyzed for each New York Bight station and survey. The

depth, grain size, carbon and nitrogen data are from summer 1980. TOe = total organic carbon; TKN = total Kjeldahl nitrogen.

--_.._.__._--

Mean

Latitude Longitude Depth Grain Size TOe TKN Months sampled (number ofsamples) - Station (ON) (OW) (m) (phi units) (mg/g dry wt.) 1979 1980 1981 1982 1983 1984 1985

1 40°26.9' 73°48.1 ' 27 3.71 9.4 1.1 Jul (1) Aug (1) Sep (2) .luI (2) Aug (2) Oct (2)

2 40°28.2' 73°45.8' 29 3.35 10.1 1.4 Jul (I) Aug (I) Sep (2) .lui (2) Aug (2) Sep (2)

Aug (2)

3 40°28.0' 73°43.8' 28 3.43 7.1 0.83 Jul (I) Aug (1) Sep (2) Jul (2) Aug (2) Oct (2)

4a 40°25.0' 73°52.0' 22 2.01 3.0 0.26 Dec (5) .luI (4) Aug (5) Jan (5) Jul (5) Aug (5) Jun (5)

Dec (5) Sep (5) Oct (2)

5 40°24.9' 73°48.0' 35 4.19 16.0 1.9 Aug (1) Aug (I) Sep (2) Jul (2) Aug (2) Oct (2)

Aug (2)

6a,b 40°25.0' 73°46.0' 27 2.73 ll.3 0.90 Dec (5) Jul (5) Aug (5) Jan (5) Jul (5) Aug (5) Jun (5)

Dec (5) Sep (5) Aug (2) Oct (2)

Nov (5) Nov (5)

7a 40°25.0' 73°44.0' 25 1.86 ~.4 0.21 Dec (5) Jul (5) Aug (5) Jan (5) Jul (5) Aug (5) Jun (5)

Dec (5) Sep (5) Nov (5) Oct (2) Q: Nov (5)

8 40°21.8' 73°51.6' 24 2.37 34.0 1.5 Jul (1) Aug (1) Sep (2) Jul (2) Aug (2) Oct (2)

Aug (2)

9 40°21.6' 73°47.8' 36 3.49 13.0 1.5 Jul (I) Aug (I) Sep (2) Jul (2) Aug (2) Oct (2) I:l:I

10 40°20.2' 73°49.1' 61 -0.24 1.6 0.17 Jul (1) Aug (I) Sep (2) Jul (2) Aug (2) Oct (2) :::

... lib 40°19.1' 73°45.8' 31 3.3 5.J 0.71 .luI (1) Aug (I) Sep (2) Jul (2) Aug (2) Oct (2) ::r

ri' 12 40°12.8' 73°44.0' 38 3.29 5.5 0.70 .luI (1) Aug (I) Sep (2) Jul (2) Aug (2) Oct (2)

13 40°09.8' 73°41.9' 56 4.25 ll.O 1.8 Jul (1) Aug (I) Sep (2) Jul (2) Aug (2) Oct (2) :-

(") 14 40°47.3' 72°59.0' 74 2.01 t.9 0.73 Aug (1) Aug (1) Sep (2) Sep (2) Aug (2) Oct (2) '"l

0 15a 40°01.4' 73°25.6' 62 3.31 8.4 1.0 Dec (5) Aug (5) Aug (5) Jan (5) Aug (5) Jun (5) ...,

:- Dec (5) Aug (5) Oct (2) :::

16 40°07.6' 73°36.8' 71 4.68 15.0 2.2 Aug (I) Aug (II Sep (2) Jul (2) Aug (2) Oct (2) 0 Sep (2) ..., 17 40°05.4' 73°31.3' 73 3.62 11.0 1.5 Aug (1) Aug (I) Sep (2) .lui (2) Aug (2) Oct (2) ;.

Sep (2) Z 18 40°25.0' 73°53.8' 24 2.2 14.0 1.1 .lui (1) Aug (1) Sep (2) Jul (2) Aug (2) Oct (2)

19 40°16.0' 73°57.8' 14 1.76 0.86 0.15 .lui (I) Aug (I) Sep (2) Jul (2) Aug (2) Oct (2)

20 40°27.9' 73°56.0' 12 1.96 0.95 0.17 Jul (I) Aug (1) Sep (2) Jul (2) Aug (2) Sep (2) ''I

21 40°31.1' 73°45.9' 21 3.15 12.0 1.5 .luI (1) Aug (I) Sep (2) Jul (2) Aug (2) Sep (2) I:l:I 22 40°25.0' 73°39.8' 24 3.07 2.4 0.36 Jul (1) Aug (1) Sep (2) Aug (2) Aug (2) Oct (2) ~.

23 39°29.9' 74°10.1 ' 16 1.01 0.46 0.09 Aug (1) Aug (I) Sep (2) Sep (2) Aug (2) Oct (2) f:

24 39°55.0' 73°55.8' 18 -0.52 1.8 0.12 Aug (I) Aug (1) Sep (2) Jul (2) Aug (2) Oct (2)

Sep (2) -

"'" 25 40°01.9' 73°55.1 ' 19 0.26 0.49 0.08 Aug (I) Aug (I) Sep (2) Jul (2) Aug (2) Oct (2) I

00 Sep (2)

26a 39°35.8' 73°54.2' 28 1.29 1.03 0.08 Dec (5) Aug (5) Aug (5) Jan (5) Jun (5) Aug (2) Jun (5)

Dec (5) Sep (5) Oct (2)
Table I (continued)

Mean Z

Grain Size TOe TKN Months sampled (number ofsamples) 0

Latitude Longitude Depth

Station (ON) (OW) (m) (phi units) (mg/g dry wt.) 1979 1980 1981 1982 1983 1984 1985

t'l

27 39°44.7' 73°44.9' 26 1.21 0.58 0.10 Aug (1) Aug (1) Sep (2) Sep (2) Aug (2) Oct (2) [ 28 39°25.5' 73°30.6' 40 2.27 0.82 0.13 Aug (1) Aug (1) Sep (2) Sep (2) Aug (2) Oct (2) t'l

29 40°14.3' 73°15.9' 38 1.09 0.63 0.13 Jul (1) Aug (1) Sep (2) Sep (2) Aug (2) Oct (2) e:-

30 40°14.8' 73°25.1' 34 1.83 0.81 0.16 Jul (1) Aug (1) Sep (2) Sep (2) Aug (2) Oct (2) ::r:;I

31 G 40°25.6' 73°11.1 ' 30 1.43 0.71 0.11 Dec \4) Jul (5) Aug (5) Jan (5) Jun (5) Aug (5) Jun (5) "0

0

Dec (5) Aug (5) Oct (2) .,

32 40°24.2' 72°58.3' 38 1.19 0.51 0.14 Jul (1) Aug (1) Sep (2) Sep (2) Aug (2) Oct (2) Z

33 40°34.1 ' 72°37.8' 40 2.37 1.8 0.29 Jul (1) Aug (1) Sep (2) Sep (2) Aug (2) Sep (2)

34 40°25.8' 72°19.8' 54 1.77 3.4 0.46 Jul (1) Aug (I) Sep (2) Sep (2) Aug (2) Oct (2) 'Jl

35 40°30.2' 72e I3.3' 55 2.34 3.9 0.69 Jul (1) Aug (1) Sep (2) Jul (2) Aug (2) Oct (2) 0- Sep (2) (.>:l

36 40°08.1' 72°51.6' 56 1.19 0.81 0.15 Aug (1) Aug (I) Sep (2) Sep (2) Aug (2) Oct (2)

37 40°04.9' 72°50.2' 54 1.41 1.0 0.17 Aug (I) Aug (1) Sep (2) Sep (2) Aug (2) Oct (2)

38 40°10.7' 72°40.3' 56 1.05 1.1 0.70 Aug (I) Aug (1) Sep (2) Sep (2) Aug (2) Oct (2)

39 40°14.'\' 73°02.0' 42 1.42 0.79 0.12 Jul (1) t.ug (1) Sep (2) Sep (2) Aug (2) Oct (2)

40 40°24.9' 73°49.7' 29 3.35 12.0 1.5 Jul (5) Aug (5) Sep (2) .lui (2) Aug (2) Oct (2)

41 40°25.0' 73°56.8' 21 1.25 0.3 0.05 Jul (5) Aug (5) Sep (2) .luI (2) Aug (2) Oct (2)

42 40°21.2' 73°56.6' 13 1.98 3.4 0.36 Aug (5) Aug (5) Sep (2) Jul (2) Aug (2) Oct (2)

43 40°18.9' 73°53.7' 20 -0.19 1.1 0.12 .lui (5) Aug (5) Scp (2) Jul (2) Aug (2) Oct (2)

44 40°13.0' 73°57.8' 19 -0.2 2.6 0.17 Jul (5) Aug (5) Sep (2) .lui (2) Aug (2) Oct (2)

63 40°39.5' 73°00.0' 12 1.04 O.H 002 Aug (1) Sep (2) Sep (2) Aug (2) Sep (2)

64 40°35.5' 73°22.0' 11 2.28 0.5 0.02 Aug (1) Sep (2) Jul (2) Aug (2) Sep (2)

65 40°3'\2' 73°37.5' 11 -0.61 0.9 0.03 Aug (1) Sep (2) Jul (2) Aug (2) Sep (2)

158 39°46.2' 73°08.2' 48 0.86 1.3 0.03 Aug (1) Sep (2) Sep (2) Aug (2) Oct (2)

i GNortheast MonitOring Program INEMP) stations.

I bAlso three samples inJul)" August, and September 1986 and 1987.

L __
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 5

because these require more data points, regularly

spaced over time.

Q-mode cluster analysis (clustering stations according to abundances of species they have in common)

was performed using all samples from the summers of

1980-85 to analyze trends in species composition

over space and time. Where multiple samples were

taken at a station and date, data were averaged. All

data were transformed by natural logarithms (In+1)

before clustering. To facilitate computation, all species occurring at <10% of the station/date combinations were eliminated. The Bray-Curtis (1957) coefficient, Cz = 2w/(a+b) , was used to measure percent

faunal similarities between stations. Here "a" is the

sum of abundances of all species found in a given

sample, "b" is the sum of species abundances for another sample, and "w" is the sum of the lower of the

abundance values for each species common to both

samples. Clustering was performed using flexible

sorting with {3 (the cluster in tensi ty Goefficien t)

= -0.25.

The Bight was divided into subareas based on geographical and depth considerations (Fig. 1) for analysis and presentation of results. The bathymetric depression at the head of Hudson Shelf Valley (HSV) is

termed the Christiaensen Basin (CB) (Fig. 2). The

Basin is actually part of the Shelf Valley (Freeland and

Swift 1978), but for this study is arbitrarily defined as

that portion north of lat. 400 20'N and in water

deeper than 27.4 m (90 feet). The shelf was divided

into NewJersey (NJ) and Long Island (LI) sections by

a line from the northwest edge of the Basin to the

middle of the Hudson-Raritan estuary's mouth. Shelf

stations were further separated into "inshore" (I: <30

m depth) and "offshore" (0: ~30 m).

Results and Discussion _

Taxa Collected

A total of 699 taxa were collected. Polychaetes were

the category with the most taxa (46% of the total),

followed by crustaceans (24%), bivalves (11 %), gastropods (9%), echinoderms (4%), coelenterates

(2%), and miscellaneous taxa (4%). The species list is

available on request.

Numbers of Species

Number of species (5) per sample is a relatively clear

indicator of environmental stress (Green 1977;

Chapman et al. 1987). Within a habitat (e.g. shallow

and sandy; deep and muddy), S is generally lower in

areas of natural or man-made stresses, whereas variables such as faunal density and diversity respond less

predictably and are more likely to increase under

moderate stress.

Spatial Trends in Numbers of Species

There were clear spatial trends in S. Figure 3 shows

contours of S values for the most recent (1985) overall survey. The lowest S by far (x = 5 species/O.l m2)

occurred at station 6, in the sludge accumulation

area. Next lowest S values were found at stations 7

(the sludge dumpsite itself), 64 (7 km SW of Fire

Island Inlet, Long Island), and 26 (26 km E of central

New Jersey) (Fig. 1). Most remaining Christiaensen

Basin stations and inshore (~30 m) stations along

both coasts had between 20 and 30 species per 0.1 m2•

Most offshore (~30 m) shelf stations had values

>30. There were trends toward increasing S with

depth as well as toward the east. Numbers of species

also increased with depth and distance from the Basin in the Hudson Shelf Valley.

Spatial trends in S were consistent among years. In

1980 (Fig. 4) as in 1985, the sludge accumulation

area had the lowest value (15/0.1 m2), followed by

Christiaensen Basin and inshore stations. Values

again increased fairly regularly from inshore to offshore and west to east on the shelf, and with depth in

the Shelf Valley.

Temporal Trends in Numbers of Species

In contrast to the consistency of spatial trends in S,

actual values of S tended to decrease over time at

many stations. Whereas much of the inshore and Basin area had S values between 30 and 40 in 1980,

almost all that area had 20-30 species/O.l m2 in 1985.

Several offshore and Shelf Valley stations had 70-80+

species in 1980, but no station had more than 63 species in 1985.

As noted in the between-year comparison of S contours, there were tendencies toward decreasing S over

time at most stations. Data for each of the six NEMP

stations, for which the most data are available, are

shown in Figure 5. Linear regression analysis of the

species versus time data for all stations indicated nonsignificant (P> 0.05) increases in S at only six of the

48 stations (Table 2). Conversely, six of the 42 decreases were significant (P"", 0.05): at stations 2, 3,
6 NOAA Technical Report NMFS 103

Table 2

Signs and significance levels of linear regression slopes fitted for numbers of species and amphipods over time (- =

decreasing, + = increasing; NS = not significant at 0.05 level). All stations combined: decreasing trends in numbers of

species (significan tat P = 0.0007) and amphipods (P = 0.0004).

Species Amphipods

Station Slope

Significance

Level Slope

Significance

Levei

New Jersey Inshore

4

8

18

19

20

23

24

25

26

27

41

42

43

44

New Jersey Offshore

28

158

Hudson Shelf Valley

11

12

13

14

15

16

17

+

NS

NS

NS

NS

NS

NS

NS

NS

NS

NS

S

NS

NS

NS

NS

NS

NS

NS

NS

0.0001

NS

NS

+

+

+

+

+

NS

NS

NS

S

NS

NS

NS

NS

NS

NS

NS

NS

0.02

NS

IS

NS

-0.002

NS

NS

0.02

0.02

0.0001

NS

0.01

Long Island Inshore

2

3

7

21

22

63

64

65

Long Island Offshore

29

30

31

32

33

34

35

36

37

38

39

+

+

0.024

0.011

0.009

NS

NS

S

NS

NS

NS

NS

NS

NS

NS

NS

NS

NS

NS

NS

NS

+

+

+

0.02

NS

NS

NS

0.03

NS

S

NS

0.01

S

NS

0.006

NS

NS

0.007

NS

NS

NS

0.005

NS

NS

0.0001

NS

0.023

NS

Christiaensen Basin

1

5

6

9

10

40

NS

NS

0.0001

0.05

l- _N~__j
Reid et a1.: Benthic Macrofauna of the New York Bight, 1979-89 7

and 7 (Long Island Inshore), 6 and 10 (Basin), and

15 (Shelf Valley). For all stations combined, there was

a decreasing trend significant at the P = 0.0007 level.

Similar trends in numbers of amphipods are discussed below.

Causes for the faunal changes are unclear. Locations of the stations with significant decreases circum-

,tantially suggest that the declines could have been

linked to increasing effects of sewage sludge with

time. (Station 6 is the presumed center of sludge accumulation, and 7 is in the dumpsite itself. Stations 2,

3, and 10 are all within 10 km of station 6. Station 15

is 52 km SE of 6, but was considered by Boesch

[1982] to be the inshore limit of a "pristine" benthic

fauna in the Shelf Valley, as of the late 1970s.) Disposal of sewage sludge increased from an average of

4,266,210 metric tons (t)/yr in 1973-79 to 6,634,355

t/yr in 1980-85 (Suszkowski and Santoro 1986), and

the recent inputs were the largest ever to any oceanic

sludge dumpsite (Norton and Champ 1989). However, the New York City Department of Environmental Protection (1983) reported that recen t increases

in sludge amounts had been due mostly to increased

water content, that sludge solids dumped increased

only 5% from 1973 to 1981, and that mass loadings of

most sludge contaminants decreased over that period. A comparison of 1973 and 1987 sludge loads

(HydroQual, Inc. 1989) indicates decreases, some

quite large, in loads of sludge solids, biochemical oxygen demand and heavy metals, although nutrient inputs increased; no 1973 data on organic con taminants are available for comparison.

There are of course other possible explanations for

the widespread declines in S (and amphipods). If anth.opogenic change in water 01" sediment quality, or

both, is involved, other waste sources may be partly or

wholly responsible. However, dredged material disposal in 1980-85 (x = 4,723,355 t/yr), decreased substantially from the 1973-79 average (9,124,785 t/yr)

(Suszkowski and Santoro 1986). Changes in carbon,

nutrient, and toxicant inputs from the HudsonRaritan estuary and other sources over the study period have not been documented. Trends in natural

environmental variables such as climate or predation

could also cause the observed changes.

Finally, there are several possible sources of sampling and analytical error. The "summer" surveys

were not all conducted at the same time of year. The

1985 survey, which showed the lowest numbers at

many stations, was conducted the latest in the year

(early October). Strict comparisons of October data

to those from mid- to late-summer may be invalid owing to seasonal differences in such variables as temperature, dissolved oxygen, sulfide, larval recruitment, and predation. Gray (1981) notes that it is

common for numbers of individuals and species to be

highest following recruitment peaks in the warmer

months, and then to decline to minima in winter. The

monthly samplings in summer 1986 and 1987 at stations 6 (Fig. 5) and II did show declines in S from

July to September, although, except for September

1986 at Station 11, all values were higher than in October 1985. Preliminary data for 1988 and 1989 again

indicate higher S values at stations 6 and 11 in summer than were found in October 1985, with slight

decreases between July and September. For both stations as well as the third primary station (northern

Christiaensen Basin) in the more recent study of responses to phaseout of sewage sludge disposal, there

were apparent increases in numbers of species between 1986 and 1989 (Environmental Processes Division, 1991). This may signal a reversal of the general

decreasing trend observed for 1980-85.

Systematic changes in sampling or sample processing are possible but unlikely explanations for the

trends, since there has been continuity of methods

and senior field and taxonomic personnel over the

study period. Sorting of samples and preliminary

identifications were done by two different contractors, but each contract included a quality assurance

program (resorting 10% of the samples to determine

and maintain accuracy). All identifications were confirmed by one of the authors (ABF).

Numbers of Amphipods

Amphipods are small crustaceans known to be important in the diets of some demersal fishes in the study

area, e.g., cod, haddock, red hake, yellowtail and winter flounder (Musick and Sedberry 1977; Langton

and Bowman 1980). Amphipods are also thought to

be relatively sensitive to chemical contamination; this

has been reported for members of the families

Ampeliscidae (Lee et a1. 1977; Sanders et a1. 1980)

and Phoxocephalidae (Swartz et al. 1982). Amphipod

densities do sometimes increase with moderate organic enrichment.

Spatial Trends in Numbers of Amphipods

Contours of amphipod densities for September 1985

(Fig. 6) show distinct spatial trends, which in general

match those described above for numbers of species.

Lowest numbers (almost always less than one amphi-
8 NOAA Technical Report NMFS 103

pod per 0.1 m2 ) were found in and near the

Christiaensen Basin. Densities were only slightly

higher at the New Jersey and Long Island Inshore

stations. Numbers increased fairly steadily to >100

amphipods/O.l m2 at the stations furthest offshore

and also in the outer Shelf Valley.

The spatial patterns were consistent over time.Figure 7 gives contours of amphipod densities for JulyAugust 1980. The chief difference from 1985 is that

in 1980 there were more inshore stations with densities elevated relative to those in the Basin. The spatial

relationships are otherwise quite similar between the

two years.

Temporal Trends in Numbers of Amphipods

Again in agreement with data on numbers of species,

there were marked decreases in numbers of amphipods at many stations over the study period. Linear

regressions of numbers vs. time indicated nonsignificant (P> 0.05) positive slopes at only nine of the 48

stations (Table 2). Fourteen of the 39 declines, however, were significant (P< 0.05): at stations 6 and 9

(Basin), 42 (New Jersey Inshore), 13, 14, 15, and 17

(Shelf Valley), 2 and 22 (Long Island Inshore), 29,

32, 35 and 39 (Long Island Offshore), and 158 (New

Jersey Offshore). For all stations combined, linear regression analysis indicated a decrease in numbers of

amphipods significant at P= 0.0004.

The most pronounced decreases (P = 0.0001) at

individual stations occurred at the same stations (6

and 15) as had the most significant declines in numbers of species. Station 6 was characterized by low

numbers of amphipods throughout the study period.

However, amphipods became more scarce with time,

decreasing from an average of 1.5/0.1 m2 in 1980-82

to only one occurrence in 27 grabs in 1983-85. Station 15 had been considered "pristine" based mostly

on the large numbers of amphipods typically found

there. Several early samplings revealed well over 1000

individuals/O.l m2, while more recent densities have

been much lower (Fig. 8). As with species richness,

the declines in amphipods could be due to anthropogenic or natural environmental change, or sampling/

analysis artifacts.

Species Composition (Cluster Analysis)

This analysis uses data on abundance and distribution

of all but rare species. It thus augments the analyses

of dominant species and numbers of species and amphipods. Since between-sample similarities are calculated using log-transformed abundances of numerous

taxa, the analysis is relatively insensitive to changes in

abundance of single taxa, e.g., amphipods. Changes

in species composition over space and time can serve

as a quantitative measure of effects of environmental

change.

The total of 292 station/date combinations (hereafter "samplings") yielded 27 groups at the 30% similarity level (Fig. 9). The 30% level was chosen for

consistency with past analysis of New York Bight data

(Reid et at. 1982), and because many of the groups

formed at that level appeared ecologically meaningful (e.g., contained most or all samplings for a given

station; contained stations that were spatially close

and had similar depths and sediment types). Some

groups contained few samplings and formed more

reasonable patterns when merged with other groups

at <30% similarity. The samplings occurring in each

group are listed in Table 3. Numerically dominant

species in each group are given in Table 4. Dominant

species are defined as those among the top ten in

abundance in at least 50% of the samplings in a

group; the actual percentages are indicated in Table

4. Distribution of the groups among various subareas

of the Bight is discussed below.

Sewage Sludge Dumpsite-Group I includes all samplings at Station 7, in the northwest corner of the

dumpsite. as well as the 1983 sampling from Station

43 (NJI). The assemblage at Station 7 is distinct in

part due to the abundance of the polychaetes,

Capitella spp., which are discussed in more detail in

Dominant Species section below. The consistency of

species composition over time at the sludge dumpsite

contradicts the suggestion from the trends in numbers of species and amphipods that degradation of

the benthos there has recently increased.

"Sewage Sludge Accumulation Area"-Group J contains all samplings at Station 6 except for October

1985, which was the sole sampling in adjacent Group

K Groups J and K joined at Cz = 0.03 similarity, and

they next merged with the sludge dumpsite fauna

(Group I), at Cz = -0.08. J was the only group in

which Capitella spp. were consistently the top

dominants.The group is also distinguished by the

consistent abundance of rhynchocoels (ribbon

worms).

Inner Hudson Shelf Valley-Group D consists exclusively of all samplings at stations 13, 16, and 17. These
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 9

Table 3

Samplings included in each group formed by cluster analysis.

Year Station (number ofsamples) Year Station (number ofsamples)

Group A Group H

1980 1(1) 2(1) 3(1) 18(1) 40(5) 1980

1981 1(1) 2(1) 3(1) 9(1) 10(1) 40(5) 1981 21(1) 25(1)

1982 1(2) 2(2) 3(2) 5(2) 8(2) 9(2) 10(2) 18(2) 40(2) 1982

1983 1(2) 2(2) 3(2) 5(2) 10(2) 18(2) 40(2) 1983 21(2) 44(2)

2(2) 5(2) 18(2) 1984

1984 1(2) 2(2) 9(2) 1985 11 (2) 21 (2) 22(2)

1985 Group I

Group B 1980 7(5)

1980 5(1) 8(1) 9(1) 1981 7(5)

1981 1982 7(5)

1982 5(2) 18(2) 1983 7(5) 43(2)

1983 9(2) 1984 7(5)

1984 1985 7(5)

1985 7(2)

Group C Groupj

1980 1980 6(5)

1981 1981 6(5)

1982 1982 6(5)

1983 1983 6(5)

1984 5(2) 10(2) 18(2) 6(2)

1985 1(2) 2(2) 3(2) 5(2) 9(2) 10(2) 18(2) 40(2) 1984 6(5)

Group D 1985 6(5)

1980 13(1) 16(1) 17(1) Group K

1981 13(1) 16(1) 17(1) 1980

1982 13(2) 16(2) 17(2) 1981

1983 13(2) 16(2) 17(2) 1982

16(2) 17(2) 1983

1984 13(2) 16(2) 17(2) 1984

1985 13(2) 16(2) 17(2) 1985 6(2)

Group E Group L

1980 II (1) 12(1) 1980 35(1)

1981 11 (I) 12(1) 1981 34(1) 35(1)

1982 II (2) 1982 35(2)

1983 II (2) 1983 35(2)

1984 35(2)

1985 1984 35(2)

Group F 1985

1980 4(4) 10(1) 21 (I) 42(5) Group M

1981 4(5) 8(1) 1980 14(1) 15(5)

1982 4(5) 21 (2) 43(2) 44(2) 1981 14(1) 15(5)

1983 8(2) 1982 14(2) 15(5)

8(2) 1983 14(2)

1984 21 (2) 1984 14(2) 15(5)

1985 8(2) 1985 14(2) 15(5)

Group G 15(2)

1980 22(1) Group N

1981 22(1) 1980 34(1) 36(1) 37(1) 38(1)

1982 22(2) 1981 36(1) 37(1) 38(1) 39(1)

1983 22(2) 1982 12(2) 32(2) 34(2) 36(2) 37(2) 38(2) 39(2)

1984 11 (2) 22(2) 1983 34(2)

1985 1984 34(2) 38(2)

1985
10 NOAA Technical Report NMFS 103

Table 3 (continued)

Year Station (number of samples) Year Station (number of samples)

Group 0

1980 29(1) 32(1) 39(1)

1981 29(1) 30(1) 33(1)

1982 29(2) 30(2)

1983 12(2) 29(2) 30(2) 32(2) 36(2) 37(2) 39(2)

1984 12(20 29(2) 30(2) 36(2) 37(2) 39(2)

1985

44(2)

44(2)

42(2) 43(2)

42(2) 43(2)

27(1)

27(1)

23(2)

23(2)

23(2)

23(2)

24(2)

19(2) 24(2) 25(2) 27(2) 28(2) 63(2)

24(2) 25(2)

24(2) 25(2) 27(2) 28(2)

24(20 25(2) 28(2)

GroupAA

1980

1981 63 (1) 65 (l )

1982 27(2) 65(2)

1983 65(2)

1984 63(2) 65(2)

1985 27(2) 63(2) 65(2)

Group X

1980 26(5)

1981 23(1) 26(5)

1982 26(5)

1983 26(5)

1984 26(2)

1985 26(5)

26(2)

Group Y

1980 41(5)

1981 41(5)

1982 41(2)

1983 41(2)

1984 41(2)

1985 41(2)

Group Z

1980 24(1) 25(1) 43(5) 44(5)

1981 24(1) 43(5)

1982

1983 43(1)

1984

1985

1984

1985

GroupW

1980 23(1)

1981

1982

1983

1984

1985

Group V

1980

1981

1982

1983

Group U (continued)

1984 4(5) 8(2)

1985 4(5)

4(2)

44(5)

64(2)

64(2)

64(2)

64(1)

20(2) 42(2)

19(1) 20(1)

42(5)

30(1) 31 (5)

31(5)

31 (5)

31(5)

31 (5)

31(2)

GroupT

1980

1981 20(1)

1982 19(2) 20(2) 42(2)

1983

1984 19(2) 20(2)

1985 19(2) 20(2)

GroupU

1980

1981

1982

1983 4(5)

Group P

1980

1981 32(1) 158(1)

1982

1983 158(2)

1984 32(2) 158(2)

1985 12(2) 29(2) 30(2) 32(2) 36(2) 37(2) 158(2)

GroupQ

1980 33(1)

1981

1982 33(2)

1983 38(2)

1984 33(2)

1985 33(2) 34(2) 35(2) 38(2) 39(2)

Group R

1980 28(1)

1981 28(1)

1982

1983

1984

1985

Group S

1980

1981

1982 28(2)

1983

1984

1985
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 11

Table 4

Consistently dominant species (in top 10 in numerical abundance in ~50% of samplings, or station/date combinations) for each group formed by cluster analysis. Am = amphipod; An = anthozoan; Bi = bivalve; Cu = cumacean;

Ec = echinoderm; Ga = gastropod; Ph = phoronid; Po = polychaete; Rh = rhynchocoel; Ta = tanaidacean.

umber of % of Number of % of

Group samplings Species Taxon samplings Group samplings Species Taxon samplings

A 33 Nucula proxima Bi 100 G 6 Nucula proxima Bi 100

Prionospio steenstrupi Po 85 Pitar morrhuanus Bi 100

Nephtys incisa Po 79 Tellina agilis Bi 83

Phoronis architecta Ph 76 Aricidea (Acesta) catherinae Po 83

Tharyx acutus Po 73 Spiophanes bombyx Po 67

Mediomastus ambiseta Po 70 Nephtys picta Po 67

Tharyx dorsobranchialis Po 67 Spio filicornis Po 67

Cossura longocirrata Po 67 Lumbrineris hebes Po 50

Ceriantheopsis americanus An 52 Tharyx acutus Po 50

B 6 Nucula proxima Bi 100 H 7 Nucula proxima Bi 100

Cossura longocirrata Po 100 Tellina agilis Bi 86

Nephtys incisa Po 83 Nephtys picta Po 86

Phoronis architecta Ph 83 Aricidea (Acesta) catherinae Po 57

Ceriantheopsis americanus An 83 Pitar morrhuanus Bi 57

Levinsenia gracilis Po 67 8 Spiophanes bombyx Po 100 Mytilus edulis Bi 67 Capitella spp. Po 88 Tharyx acutus Po 50 Tharyx acutus Po 88 Pherusa affinis Po 50 Nephtys picta Po 88

C 11 Ceriantheopsis americanus An 100 Tellina agilis Bi 88

Nucula proxima Bi 100 Exogone hebes Po 63

Nephtys incisa Po 100 Nephtyidae spp. Uuvenile) Po 50

Pherusa affinis Po 82 J 7 Capitella spp. Po 100 Phoronis architecta Ph 73

Cossura longocirrata Po 55 Rhynchocoela spp. Rh 100

Lumbrineris hebes Po 55 Nephtys incisa Po 86

Nucula proxima Bi 86

0 20 Nucula delphinodonta Bi 100 Spio filicornis Po 71

Cossura longocirrata Po 95 Pholoe minuta Po 57

Levinsenia gracilis Po 95

Ninoe nigripes Po 85 K (one sampling only)

Tharyx dorsobranchialis Po 80 Nephtys incisa Po 100

Aricidea (Acesta) catherinae Po 75 Glycera dibranchiata Po 100

Euchone incolor Po 75 Pherusa affinis Po 100

Lumbrineris hebes Po 70 Nassarius lrivit/alus Ca 100

Prionospio steenstrupi Po 70 Nucula proxima Bi 100

Mediomastus ambiseta Po 50 Tellina agilis Bi 100

E 6 Nucula proxima Bi 100 Spisula solidissima Bi 100

Rhynchocoela spp. Rh 83 L 7 Ampelisca agassizi Am 100

Tharyx dorsobranchialis Po 83 Unciola irrorata Am 100

Ninoe nigripes Po 83 Leptocheirus pinguis Am 100

Aricidea (Acesta) catherinae Po 83 Corophium crassicorne Am 100

Lumbrineris hebes Po 83 Erichthonius rubricornis Am 86

Prionospio steenstrupi Po 83 Unicola spp. Uuvenile) Am 71

Spio filicornis Po 50 Exogone verugera Po 71

Pholoe minuta Po 50 Harpinia propinquua Am 57

F 14 Amastigos caperatus Po 79 Eudorella pusilla Cu 57

Tellina agilis Bi 79 M 12 Ampelisca agassizi Am 100

Mediomastus ambiseta Po 71 Aricidea (Acesta) catherinae Po 83

Nucula proxima Bi 64 Lumbrineris hebes Po 75

Aricidea (Acesta) catherinae Po 64 Unciola irrorata Am 58

Tharyx acutus Po 57 Tharyx dorsobranchialis Po 50

Parougia caeca Po 50 Nunda delphinodonta Bi 50

Prionospio steenstrupi Po 50 Leptocheirus pinguis Am 50
12 NOAA Technical Report NMFS 103

Table 4 (continued)

umber of % of i Number of % of

Croup samplings Species Taxon samplings .-1 Group samplings Species Taxon samplings

N 18 Exogone hebes Po 100 T (continued) Pseudunciola obliquua Am 58

Unciola inermis Am 78 Spiophanes bomilyx 1;'0 50

Byblis serrata Am 72 Nephtyidae spp. (juvenile) Po 50

Aricidea (Acesta) catherinae Po 61 Protohaustorius cf

Caulleriella cf killariensis Po 56 deichmannae Po 50

Euchone elegans Po 56 LJ 17 Tellina agilis Bi 88

0 21 Unicola inermis Am '00 Goniadella gracilis Po 64

Exogone hebes Po Y5 Spiophanes bomilyx Po 64

Caulleriella cf killariensis Po 90 Spisula solidissima Bi 64

Tanaissus liljeborgi Ta 67 Caulleriella cf killariensis Po 59

Rhepoxynius hudsoni Am 57 lephtyidae spp. (juvenile) Po 59

Prionospio steenstrupi Po 52 V 18 Echmarachnius parma Ec 78

Aricidea (Acesta) catherinae Po 52 Goniadella gracilis Po 72

Byblis serrata Am 5~ Astarte castanea Bi 67

P 12 Caulleriella cf killariensis Po 100 Exogone hebes Po 61

Exogone hebes Po 100 Caulleriella if. killariensis Po 61

Goniadella gracilis Po 100 Pseudunciola obliquua Am 61

Aricidea (Acesta) catherinae Po 83 Tanaissus liljeborgi Ta 50

Lumbrinerides acuta Po 75 W 5 Goniadella gracilis Po 100

Nephtydae spp. (juvenile) Po 58 Tellina agilis Bi 80

Unciola irrorata Am 58 Echinarachnius panna Ec 80

'J'anaissus liljeborgi Ta 50 Pseudunciola obliquua Am 80

Q 9 Exogone hebes Po 56 Astarte castanea Bi 60

Rhepoxynius hudsoni Am 56 Spisula solidissima Bi 60

Unciola iner mis Am 56 Hemipodus roseus Po 60

Ampelisca agassizi Am 56 Sigalion arenicola Po 60

R 2 Phyllodoce mucosa Po 100 Nephtys bucera Po 60

Glycera spp. (juvenile) Po 50 Rhynchocoela spp. Rh 60

Exogone hebes Po 50 X 8 Echinarachnius parma Ec 100

Aricidea (Aricidea) wassi Po 50 Ceriantheopsis americana An 88

Corophium crassicorne Am 50 Tharyx dorsobranchialis Po 88

Aglaophamus circinati Po 50 Nephtyidae spp. (juvenile) Po 75

Uncio/a inermis Am 50 Goniadella gracilis Po 63

Paraonis Julgens Po 50 Nephtys picta Po 63

Echinarachnius parma Ec 50 Tharyx acutus Po 63

Spiophanes bombyx Po 50 Hemipodus roseus Po 50

Parougia caeca Po 50 Lumbrineris acicularum Po 50

Clymenella torquata Po 50 y 6 Parapionosyllis longicirrata Po 100

Cerastoderma pinnulatum Bi 50 Goniadella gracilis Po 100

Lumbrineris hebes Po 50 Spisula solidissima Bi 100

Rhepoxynius hudsoni Am 50 Tanaissus liljeborgi Ta 100

Sthenelais limicola Po 50 Tellina agilis Bi 83

Photis macrocoxa Am 50 Aricidea (Acesta) cerruti Po 67

Rhynchocoela spp. Rh 50 Nephtys bucera Po 50

Ceriantheopsis americans Am 50 ephtyidae spp. (juvenile) Po 50

S 9 Exogone hebes Po 89 Scolelepis squamata Po 50

Caulleriella cf killariensis Po 89 Spiophanes bomilyx Po 50

Pseudunciola obliquua Am 89 Hemipodus roseus Po 50

Aricidea (Acesta) catherinae Po 67 Z 7 Aricidea (Acesta) catherinae Po 100

Tanaissus liljeborgi Ta 67 Lumbrineris acicularum Po 100

Rhepoxynius hudsoni Am 56 Goniadella gracilis Po 86

Echinarachnius parma Ec 56 Parougia caeca Po 86

T 12 Tellina agilis Bi 92 Tharyx acutus Po 71

Nephtys picta Po 75 AA 10 Cirrophorus brevicirratus Po 100

Spisula solidissima Bi 67 Goniadella gracilis Po 90

Tanaissus liljeborgi Ta fj7 Hemipodus roseus Po 80

Aricidea (Acesta) catherinae Po 58 Aricidea (Acesta) catherinae Po 80

Caulleriella cf killariensis Po 58 Tellina agilis Bi 50

Magelona riojai Po 58
Reid et aI.: Benthic Macrofauna of the New York Bight, 1979-89 13

stations are located in the inner HSV, 28-41 km SSE

of the sludge dumpsite and sludge accumulation

area. That all 1980-85 samplings from the three stations clustered together, and without any trend toward early samplings segregating from later ones, is

evidence against any gross effects of increasing sludge

disposal or other environmental changes in the inner

HSV over that period. However, the preponderance

of polychaetes among the dominant species may be

an indication of long-term enrichment or stress effects. Group D clustered most closely with Groups A,

B, and C (samplings from the Christia~nsen Basin

and stations closest to the Hudson-Raritan estuary

mouth), another sign that influences in those areas

extend to the inner HSV.

The other two stations in the inner HSV clustered

separately from stations 13, 16, and 17 (Group D).

Station 11, just south of the Christiaensen Basin, was

in the adjacent though quite dissimilar (Cz = -1.4)

Group E for 1980 through 1983, and St£\tion 12, located between 11 and 13, was in E in 1980 and 1981.

Station 11 moved to Group G in 1984 and H in 1985;

it thus moved closer to the sludge dumpsite and accumulation area on the dendrogram but was still quite

dissimilar from them (Cz = -0.36). Station 12 samplings from 1982 through 1985 fell in the "offshore

supergroup," with amphipods increasing in abundance in contrast to the trend at most stations.

Outer Hudson Shelf Valley-Unlike the inner HSV

and CB, stations 14 and 15 (Group M) were dominated by amphipods. As with Group D in the inner

HSV, the similarity (0.45) of all samplings in the

outer HSV is evidence against major effects of environmental change over the study period, and contradicts the decreases in numbers of species and amphipods discussed above.

Long Island Offshore (LIO)-The fauna of the easternmost LIO stations was most similar to that of the

outer HSV. Group L, which contained samplings from

stations 34 and 35, joined the outer HSV group at

0.15 similarity. As in the outer HSV, this group was

dominated by Ampelisca agassizi and other amphipods,

and had a relative scarcity of polychaetes. A. agassizi

did decrease somewhat from the very high 1980-81

levels, and then increased again in 1985 when Station

35 fell into Group Q (discussed below). Polychaetes

were more dominant in Group Q, so the"re are mixed

indications of temporal change in species composition in this area.

All other LIO stations had somewhat similar faunas.

All samplings fell within groups N, 0, P, Q, and S,

and all these groups joined at -0.21 similarity. The

largest difference between these groups and Group L

above was their lower abundance of Ampelisca agassizi.

New Jersey Offshore-The two stations (28 and 158)

in this category had assemblages most similar to LIO

stations. All samplings from Station 158 fell in Group

P, which otherwise consisted of LIO samplings. The

1980 and 1981 samplings at Station 28 formed a separate group (R), with closest affinities to LIO Station

31. Station 28 occurred in Group S in 1982, and

Group V (New Jersey ·Inshore, below) in 1983

through 1985.

New Jersey Inshore-This is the largest (14 stations),

most heterogeneous collection of stations. Samplings

were split between two very dissimilar (Cz = -3.0)

"supergroups," A through K and T through AA. The

NJI stations closest to the Christiaensen Basin (4, 8,

and 18) had most samplings in the A-K supergroup.

Groups T through AA basically consisted of samplings

from the more southern and inshore NJI stations (19,

20, 23-27 and 41-44). The cluster analysis did not

distinguish northern coastal stations from other NJI

areas, indicating that the Hudson-Raritan estuarine

plume (which tends to follow the north NJ coast)

did not have an overriding influence on species

composition.

Long Island Inshore-As with NJI stations, the LII

stations were split between the dissimilar supergroups

A-K and T-AA. All samplings from the three stations

nearest the Long Island coast (63, 64, and 65) were in

group T-AA. All sampling from the remaining LII

stations (2, 3, 21, and 22) were in groups A, C, F, G,

and H, with fauna quite distinct from stations 63-65.

Christiaensen Basin-Finally, all remaining CB stations (1, 5, 9, 10, and 40) clustered together in

groups A-C, except for Station 10 in 1980 (Group F).

These are the stations physically closest to Station 6 in

the sludge accumulation area. Their depths are similar to that at Station 6 (Table 1) and, as indicated by

amounts of fine sediments, they also have somewhat

depositional environments (Station 10 is deeper but

has coarser sediments). These stations should be

among the first to show signs of any increasing influence of sludge or other contaminant inputs to the

Basin. That their species composition remained consistent over time, and distinct from the fauna at Station 6, from 1980-85 is evidence against large increases in anthropogenic effects.
14 NOAA Technical Report NMFS 103

Table 5

Mean densities and biomasses per 0.1 m with standard errors and numbers of replicate samples analyzed, for the

numerically dominant species at Station 6 on each sampling date.

No. of Density Biomass (mg.)

Date samples Species Mean SE Species Mean SE

Dec 1979 5 grabs Phoronis architecta 764.0 205.5 Phoronis architecta 2321.0 707.0

Asabellides oculata 490.8 81.2 Nephtys incisa 1688.2 474.3

Tharyx acutus 342.2 38.4 Rhynchocoela spp. 1440.4 334.4

Capitella spp. 118.6 28.3 Ceriantheopsis americanus 1248.4 380.9

Mediomastus ammseta 79.2 11.7 Pherusa affinis 467.6 254.9

Cossura longocirrata 68.2 13.1 Ninoe nigripes 375.6 245.5

Prionospio steenstrupi 50.0 12.9 Asabellides oculata 372.4 51.7

Parougia caeca 45.8 13.7 Cancer irroratus 297.6 251.0

Unciola irrorata 44.6 7.2 Crangon septemspinosa 106.4 38.0

Nephtys incisa 42.4 8.2 Tharyx acutus 82.6 5.9

Jul 1980 5 grabs Capitella spp. 545.6 118.4 Capitella spp. 497.2 245.5

Edotea triloba 27.4 8.5 Rhynchocoela spp. 201.0 109.3

Rhynchocoela spp. 8.4 3.0 Edotea triloba 111.2 38.3

Spiophanes bomlJyx 8.2 6.8 Ceriantheopsis americanus 106.4 93.4

Cancer irroratus 7.0 3.4 Cancer irroratus 70.8 46.0

Tellina agilis 4.2 2.1 Spiophanes bomlJyx 56.4 50.6

Tharyx acutus 3.6 2.7 Nephtys incisa 42.4 18.3

Nephtys incisa 3.2 1.6 Phoronis architecta 15.4 13.9

Unciola irrorata 2.2 0.6 Pherusa affinis 15.2 13.3

Pitar morrhuanus 1.8 1.8 Tellina agilis 12.8 6.8

Dec 1980 5 grabs Spiophanes bomlJyx 166.2 34.4 Ceriantheopsis americanus 832.2 434.5

Capitella spp. 86.4 44.5 Nephtys picta 254.8 67.3

Pherusa affinis 46.0 14.5 Pherusa affinis 228.8 79.5

Nephtyidae spp. 35.8 6.9 Diopatra cuprea 173.6 132.8

Nephtys pieta 14.2 2.8 Spiophanes bomlJyx 167.6 18.6

Parougia caeca 11.0 4.0 Glycera dibranchiata 62.6 23.3

Edotea tribola 5.4 1.7 Capitella spp. 49.8 29.7

Tharyx acutus 4.8 2.5 Rhynchocoela spp. 45.8 45.1

Ceriantheopsis americanus 3.4 1.7 Pitar morrhuanus 15.2 15.2

Diopatra cuprea 3.4 2.0 Edotea tribola 13.4 5.3

Aug 1981 5 grabs Capitella spp. 637.8 157.7 .Capitella spp. 2475.6 630.9

Rhynchocoela spp. 27.2 6.8 Rhynchocoela spp. 1670.4 509.2

Phoronis architecta 14.6 6.3 Nephtys incisa 223.8 39.8

Nucula proxima 7.4 2.6 Nucula proxima 82.6 26.8

Phyllodoce (anatides) mucosa 7.0 2.1 Ovalipes ocellatus 49.4 49.4

Nephtys incisa 6.4 0.6 Cancer irroratus 28.6 14.6

Pholoe minuta 3.8 0.4 Phersusa affinis 22.8 12.3

Microphthalmus sczelkowii 2.0 0.8 Phoronis architecta 19.2 8.4

Prionospio steenstrupi 2.0 1.4 Tharyx dorsobranchialis 17.6 17.1

Tharyx acutus 2.0 0.9 Ceriantheopsis americanus 14.2 9.3

Jan 1982 5 grabs Nucula proxima 51.4 13.7 Rhynchocoela spp. 337.2 107.3

Phoronis architecta 16.2 7.3 Nucula proxima 270.0 55.3

Edotea triloba 4.4 2.0 Nephtys incisa 189.0 60.3

Rhynchocoela spp. 3.8 0.6 Phersusa affinis 49.6 36.7

Tharyx acutus 3.8 1.6 Phoronis architecta 40.2 16.8

Nephtys incisa 3.6 1.2 Edotea triloba 11.6 4.4

Capitella spp. 3.2 0.9 Amphioplus abditus 8.2 8.2

Nephtyidae spp. 2.4 1.2 Spiophanes bomlJyx 5.8 2.2

Cirratulus cirratus 2.4 2.4 Ceriantheopsis americanus 5.2 3.6

Parougia caeca ~.2 1.1 Crangon septemspinosa 5.2 4.5
Reid et aI.: Benthic Macrofauna of the New York Bight, 1979-89 15

Table 5 (continued)

No. of Density Biomass (mg)

Date samples Species Mean SE Species Mean SE

Aug 1982 5 grabs Capitella spp. 989.2 379.0 Rhynchocoela spp. 2139.4 486.7

Ceriantheopsis americanus 55.2 18.6 Capitella spp. 360.0 107.5

Pholoe minuta 41.4 16.9 Ceriantheopsis americanus 334.8 76.9

Spio filicornis 16.6 4.4 Nephtys incisa 316.0 95.3

Phyllodoce (anatides) mucosa 16.2 4.3 Cancer irroratus 162.6 100.0

Rhynchocoela spp. 14.4 3.2 Spio filicornis 140.6 41.5

Parougia caeca 9.0 1.8 Pherusa affinis 18.2 17.5

Tharyx acutus 8.4 3.4 Phyllodoce (anatides) mucosa 17.0 4.5

Nucula proxima 8.4 1.8 Crangon septemspinosa 17.0 17.0

Mediomastus ambiseta 7.4 3.8 Glycera dibranchiata 16.2 13.6

Nov 1982 5 grabs Nucula proxima 11.8 4.7 Nephtys incisa 364.0 103.5

Nephtys incisa 8.6 1.9 Ceriantheopsis americanus 30.6 30.6

Diastylis polita 8.2 2.6 Rhynchocoela spp. 28.2 19.9

Pherusa affinis 6.6 2.8 Nucula proxima 25.2 10.2

Tellina agilis 5.8 3.0 Pherusa affinis 19.2 7.8

Asabellides oculata 2.2 1.2 Diastylis polita 10.8 3.7

Spiophanes bombyx 2.0 0.9 Crangon septemspinosa 10.8 4.7

Edotea tribola 1.8 0.6 Asabellides oculata 5.8 5.1

Crangon septemspinosa 1.6 0.7 Edotea triloba 2.4 0.9

Capitella spp. 1.4 0.7 Spiophanes bombyx 2.2 1.5

Jul 1983 5 grabs Capitella spp 86.8 52.5 Capitella spp. 358.0 125.2

Rhynchocoela spp. 4.0 1.8 Rhynchocoela spp. 119.0 71.0

Pholoe minuta 3.2 2.2 Nucula proxima 18.8 18.1

Nucula proxima 2.6 1.9 Nephtys incisa 12.4 9.8

Tharyx dorsobranchialis 1.6 1.6 Spio filicornis 2.8 2.6

Nephtys incisa 1.4 0.7 Harmothoe extenuata 2.0 2.0

Spio filicornis 1.4 0.9 Pholoe minuta 0.8 0.6

Ceriantheopsis americanus 0.4 0.4 Tharyx dorsobranchialis 0.8 0.8

Mytilus edulis 0.4 0.4 Cancer irroratus 0.8 0.8

Harmothoe extenuata 0.2 0.2 Ceriantheopsis americanus 0.6 0.6

Aug 1983 2 grabs Capitella spp. 5604.0 5092.0 Capitella spp. 3985.5 3643.5

Rhynchocoela spp. 39.5 6.5 Rhynchocoela spp. 959.0 200.0

Ceriantheopsis americanus 23.5 15.5 Ceriantheopsis americanus 212.5 188.5

Pholoe minuta 16.0 7.0 Nucula proxima 46.0 4.0

Nucula proxima 14.5 1.5 Spio filicornis 12.0 3.0

Phoronis architecta 4.0 3.0 Nephtys incisa 10.0 0.0

Nephtys incisa 3.0 0.0 Pherusa affinis 5.0 5.0

Spio filicornis 3.0 0.0 Edwardsia elegans 2.5 2.5

Parougia caeca 2.0 1.0 Pholoe minuta 2.5 0.5

Prionospio steenstrupi 2.0 1.0 Eteone longa 2.0 2.0

ov 1983 5 grabs Capitella spp. 414.8 368.8 Nephtys incisa 1082.4 305.0

Nereis succinea 35.2 31.7 Pherusa affinis 321.8 241.9

Nucula proxima 28.2 6.6 Rhynchocoela spp. 296.0 85.6

Neplttys incisa 27.2 5.2 Capitella spp. 191.4 175.5

Parougia caeca 21.0 9.8 Ceriantheopsis americanus 181.8 44.5

Asabellides oculata 16.2 3.5 Nucula proxima 120.0 28.8

Nephtys picta 12.0 4.1 Dichelopandalus leptocerus 56.0 56.0

Pherusa affinis 4.8 2.6 Nereis succinea 55.6 54.1

Tellina agilis 4.0 1.3 Asabellides oculata 37.0 12.7

Ceriantheopsis americanus 34 0.8 Tellina agilis 6.2 2.6
16 NOAA Technical Report NMFS 103

r----- ---_._--,

I Table 5 (continued) I , Density Biornas~ (IT.g.) I

I

t\o. of

Date sa!nple~ Sp"c:e, Mean SE Species Meal~ SE

Aug 1984 5 grab~ Capitella spp. 117.4 54.2 Capitella spp. 13104 831.5

Rhynchocoela spp. 3.4 1.3 Rhynchocoela spp. 336.6 16g.~

Nucuia proxima 1.0 0.6 Tellina agilis 12 R 7.8

Pherusa affinis 0.8 0.6 Pherusa affinis ; 1.4 7.2

Nephlys incisa 0.6 0.4 Nephty incisa 7.6 5.2

Edotea lriloba 0.6 0.4 Ceriantheapsis americanu.~ 7.0 7.0

Spio fiiicornis 0.4 0.4 Crangon Septemspinosu 4.0 2.5

Tellina agilis 0.4 0.2 Nucula proxima 3.0 2.0

Crangon Septemspinosa 0.4 0.2 Edotea triioba 2.8 2.0

Ceriantheopsis americanus 0.2 0.2 Spio filicornus 1.8 1.8

Jun 1985 5 grabs Capitella spp. 99.8 45.0 Capitella spp. 326.8 193.0

Nucula p1'Oxzma 19.6 4.7 Rhynchocoela spp. 239.6 113.5

Nephtys inci,a 12.6 1.1 Nephty!> incisa 196.6 40.3

Arclic', islandica. 5.2 2.0 Nucula p1'Oxima 126.8 30.3

Edotea Inloba 3.6 1.0 Nassarius trimttatus 17.8 17.8

Rhynchvcoela spp. 2.4 0.5 Edotea Triloba 11.2 1.7

Anthozoa spp. 2.2 0.7 Arlica islandica 6.0 2.7

Spio fiiicornus 1.0 0.3 Cancer irroraius 5.0 2.3

Cancer zrroralus 1.0 0.4 Mulinia laleralis 4.0 2.5

Crangon sl!"ptemspinosa 0.6 0.6 Anthozoa spp. 3.2 1.0

Oct 1985 2 grabs Nephtys incisa 7.0 1.0 Nephtys incisa 180.0 4.0

Nassarius trivillatus 1.0 0.0 Glycera dibranchiata 30.0 30.0

Nucuia proxima 1.0 1.0 Tellina agiiis 3.0 1.0

Tellina agilis 10 0.0 Nassarius trivittatus 2.5 0.5

Giycera dibranchiata 0.5 LJ.f:- Nucuia proxima 2.5 2.5

Pherusa affinis 0.5 0.5 Phnusa aJfinis 1.0 1.0

Spisuia soiidissima 0.5 0.5 Spzsuia solidissima 1.0 1.0

Jul1986 3 grabs Capitella spp. 2177.3 1433.3 Rhynchocoela spp. 3792.3 1478.7

Asabellides oculata 486.3 92.7 Asabellzdes oculata 3128.0 1440.7

Pherusa affinis 72.3 39.1 Capitella spp. 2552.7 2289.1

Nucuia p1'Oxima 68.0 42.0 Pherusa affinis 622.0 365.2

Rhynchocoela spp. 60.3 17.9 Paranaitis speciosa 287.0 26.2

Tellina agilis 50.3 33.2 Nephlys incisa 253.3 237.0

Paraougia caeca 40.0 40.0 Ceriantheopsis americanus 243.7 217.8

Paranailus sp~ciosa 31.7 3.2 Cancer irroratus 219.0 81.2

Thm yx acutus 24.0 10.0 Asteriasforbesi 122.7 122.7

Cancer irroratus 16.3 7.4 Nucula proxima 119.7 48.0

Aug 1986 3 grabs Tharyx spp. 33.7 33.7 Rhynchocoela spp. 2283.0 835.0

Rhynchocoela spp. 23.3 11.6 Cancer i,-,-oratus 891.0 611.3

Nucuia proxima 22.0 18.0 Nephtys incisa 521.0 278.9

Tharyx acuttH 16.7 14.7 Ceriantheopsis amaicanus 361.0 209.3

Canur irroratus 12.3 2.6 Nucula proxima 63.0 41.7

Cerioantheopsis americanus 12.0 8.3 Pherusa affinis 30.3 13.0

Pherusa affinis 11.3 6.4 Nassarius trivittatus 28.0 28.0

Nephtys incisa 4.7 2.3 Tha,-yx spp. 7.3 7.3

Mediomastus ambisp,/a 4.3 2.3 Thar)'x .:lcutus 7.3 5.9

Tellina agilis 2.3 1.5 PhoTOnis architecta 7.0 6.0

Sep 1986 3 grabs Rhynchocoela spp. 39.7 14.4 Rhynchocoela spp. 2169.0 853.0

Tharyx act/Ius 22.3 5.9 Cerianlheapsis ame'licanus 552.0 251.0

Nucula proxima 22.0 10.2 Pherusa affinis 199.7 193.2

Cerianlhcop,is americanus 20.3 9.0 Nephtys incisa 135.0 45.3
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-8£ 17

Table 5 (continued)

o. of Density Biomass (mg)

Date samples Species Mean SE Species Mean SE

Capitella spp. 6.7 3.3 Glycera dibranchiata 94.3 52.6

Mediomastus ambiseta 6.3 2.3 Nucula proxima 51.3 15.8

Nephtys incisa 2.3 0.9 Spio fiiicornis 11.0 11.0

Tellina agilis 2.3 1.5 Ninoe nigripes 8.0 8.0

Parougis caeca 2.0 2.0 Tharyx acutus 8.0 3.6

Pherusa a/finis 2.0 1.2 Tellina agilis 2.3 1.9

Jul 1987 3 grabs Rhynchocoela spp. 71.7 40.2 Rhynchocoela spp. 9054.7 4015.4

Parougia caeca 37.0 21.0 Ceriantheopsis americanus 1507.0 796.7

Cenantheopsis americanus 36.3 21.3 Nephtys incisa 243.3 224.4

Nucula proxima 27.3 13.7 Nucula proxima 134.7 58.7

Capitella spp. 15.0 14.0 Edotea tnloba 106.0 36.8

Edotea tnloba 13.0 3.5 Parougia caeca 45.0 28.4

Cancer irroratus 4.3 2.8 Cancer irroratus 36.3 28.3

Nephtys incisa 3.7 1.5 Pitar morrhuanus 18.3 18.3

Spio /ilicornis 2.0 2.0 Spio filicornis 10.0 10.0

Harmothoe extenuata 1.3 0.7 Spiophanes bombyx 8.7 8.7

Aug 1987 3 grabs Parougia caeca 48.3 29.1 Rhynchocoela spp. 3691.0 1441.4

Rhynchocoela spp. 37.0 16.1 Ceriantheopsis americanus 782.7 530.0

Capitella spp. '34.7 16.8 Nucula proxima 152.3 70.4

Nucula proxima 25.7 13.6 Pherusa a/finis 45.0 23.1

Ceriantheopsis amencanus 9.3 6.2 Edotea t,iloba 34.0 21.1

Edotea tnloba 5.0 3.1 Spio filicornis 22.3 22.3

Pherusa a/finis 2.0 0.6 Cancer irroratus 21.7 11.5

Cancer irroratus 1.3 0.9 Parougia caeca 15.3 10.5

Spio filicornis 1.0 1.0 Nephtys incisa 9.7 9.7

Tharyx acutus 0.7 0.7 Capitella spp. 6.7 3.8

Sep 1987 3 grabs Capitella spp. 2187.3 867.3 Rhynchocoela spp. 3542.7 1172.9

Nucula proxima 63.0 29.6 Ceriantheopsis americanus 1429.7 1336.3

Rhynchocoela spp. 22.7 5.2 Capitella spp. 1089.3 394.4

Ceriantheopsis americanus 19.0 18.0 Cancer irroratus 936.3 936.3

Parougia caeca 4.3 2.0 Nucula proxima 154.7 23.7

Pnonospio steenstrupi 3.3 2.8 Nephtys incisa 26.7 12.1

Edotea tnloba 2.7 1.2 Pherusa a/finis 17.3 17.3

Nephtys incisa 2.3 0.7 Parougia caeca 5.0 2.6

Unciola irrorata 1.0 1.0 Edotea tnloba 3.7 1.5

Jassa /alcata 0.7 0.3 Dyopedos monocanthus 3.3 1.7
18 NOAA Technical Report NMFS 103

Table 6

Mean densities and biomasses per 0.1 m with standard errors and numbers of replicate samples analyzed, for the

numerically dominant species at Station 7 on each sampling date.

No. of Density Biomass (mg)

Date samples Species Mean SE Species Mean SE

Dec 1979 5 grabs Capitella spp. 372.0 18! .5 Diopatra cuprea 1032.8 479.6

Tharyx acutus 296.0 46.1 Ceriantheopsis americanus 633.2 201.4

Parougia caeca 98.8 22.8 Nephtys picta 631.8 99.9

Exogone hebes 69.8 10.4 Rhynchocoela spp. 523.4 231.4

Spiophanes bombyx 60.8 23.9 Asabellides oculata 300.2 64.5

Asabellides oculata 57.4 24.0 Ensis directus 262.8 158.4

Unciola irrorata 44.8 20.2 Spiophanes bombyx 220.4 59.9

Mephtys picta 37.8 3.0 Nassarius trivittatus 209.4 129.5

Nephtyidae spp. 23.0 5.1 Glycera dibranchiata 180.4 22.5

Phoronis architecta 13.4 5.8 Tharyx acutus 177.2 26.3

Jul 1980 5 grabs Edotea triloba 62.0 !3.6 Lumbrineris acicularum 522.0 334.6

Capitella spp. 42.8 16.8 Ceriantheopsis americanus 351.6 260.8

Tharyx acutus 28.6 12.7 Rhynchocoela spp. 280.3 98.9

Tellina agilis 23.3 5.4 Edotea triloba 236.4 89.1

Nephtys picta 17.4 10.2 Nephtys picta 225.6 141.1

Amastigos caperatlls 15.7 9.9 Glycera dibranchiata 145.6 105.4

Parougia caeca 13.4 5.5 Tellina agilis 111.1 44.4

Spiophanes bombyx 13.4 5.9 Nephtys incisa 102.8 102.8

Tharyx acutus 11.7 11.6 Spiophanes bombyx 99.4 74.8

Exogone hebes 9.1 9.0 Nephtys bucera 73.3 73.2

Dec 1980 5 grabs Spiophanes bombyx 108.5 26.1 Arctica islandica 15400.0 15400.0

Capitella spp. 32.6 11.0 Nassarius trivittatus 1091.1 615.5

Parougia caeca 30.8 18.4 Diopatra cuprea 417.0 415.8

Pherusa a/finis 25.3 12.3 Ceriantheopsis americanus 403.5 189.0

Nephtyidae spp. 14.7 5.2 Nephtys picta 251.1 132.9

Tharyx acutus 10.1 6.1 Rhynchocoela spp. 207.9 169.8

Nephtys picta 9.1 2.9 Spiophanes bombyx 182.6 124.6

Diopatra cuprea 4.5 3.7 Pherusa a/finis 107.3 55.0

Exogone hebes 3.8 2.6 Crangon septemspinosa 28.7 4.2

Nereidae spp. (iuv.) 3.5 2.1 Laonice cirrata 17.9 17.8

Aug 1981 5 grabs Spiophanes bombyx 158.4 29.5 Spiophanes bombyx 1516.7 210.2

Capitella spp. 79.1 33.8 Diopatra cuprea 370.0 370.0

Tharyx acutus 24.6 20.7 Ceriantheopsis americanus 346.7 97.5

Exogone hebes 17.8 2.6 Rhynchocoela spp. 216.6 103.4

Spio filicornis 13.9 5.7 Nephtys pieta 187.4 44.5

Tellina agilis 13.3 6.6 Nassarius trivittatus 158.3 158.2

Tharyx dorsobranchialis 11.1 9.5 Spio filicornis 99.7 31.5

Ceriantheopsis americanus 10.8 1.1 Tellina agilis 91.7 38.0

Nephtys picta 8.9 3.2 Glycera dibranchiata 71.7 29.4

Nephtyidae spp. 4.9 3.1 Capitella spp. 60.9 17.2

Jan 1982 5 grabs Echinarachnius parma 192.4 48.5 Spiophanes bombyx 1428.6 336.2

Spiophanes bombyx 120.4 14.0 Ceriantheopsis americanus 630.0 226.1

Nephtyidae spp. 82.6 6.7 Rhynchocoela spp. 317.4 143.9

Exogone hebe~ 26.0 6.!l Nephtys picta 202.4 37.5

Tellina agilis 25.4 3.8 Diopatra cuprea 200.6 179.6

Parougia caeca 22.2 5.3 Nassarius trivittatus 121.8' 79.9

Tharyx acutus 17.8 6.2 Nephtyidae spp. 68.8 9.9

Cap~tella spp. 17.2 2.7 Pherusa a/finis 60.0 33.3

Nephtys picta 7.6 I.::? Glycera dibranchwta 53.6 20.1

Glycera spp. (iuv.) 7.2 1.9 Tellina agilis 33.4 20.8
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 19

Table 6 (continued)

No. of Density Biomass (mg)

Date samples Species Mean SE Species Mean SE

Aug 1982 5 grabs Capitella spp. 678.3 123.7 Rhynchocoela spp. 8553.3 1749.8

Parougia caeca 167.4 12.8 Ceriantheopsis amerieanus 2080.6 903.1

Exogone hebes 79.3 26.7 Capitella spp. 1915.3 356.1

Spiophanes bombyx 44.6 16.0 Spiophanes bombyx 875.7 343.7

Nephtys pieta 38.0 17.0 Nephtys pieta 533.7 222.9

Tharyx acutus 24.4 3.7 Diopatra cuprea 368.0 185.4

Tellina agilis 17.8 5.0 Nassarius triviUatus 365.6 218.5

Rhynchocoela spp. 13.7 4.6 Glyeera dibranehiata 277.5 167.0

Ceriantheopsis amerieanus 5.1 1.2 Cancer irroratus 188.3 126.0

Amastigos caperatus 3.8 1.7 Tellina agilis 164.1 76.6

Nov 1982 5 grabs Capitella spp. 82.6 44.1 Diopatra euprea 1597.2 1294.4

Nereis suecinea 27.8 23.5 Pitar morrhu...nus 924.0 924.0

Spio filieornis 23.8 14.0 Spio filieornis 179.0 113.8

Spiophanes bombyx 14.4 6.0 Glyeera dibranehiata 168.4 119.0

Pherusa affinis 9.8 3.0 Rhynchocoela spp. 103.6 71.3

Diopatra cuprea 9.0 6.3 Ceriantheopsis americanus 88.4 63.3

Gammarus lawreneianus 8.6 8.6 Capitella spp. 83.0 46.0

Euehone ineolor 6.6 6.4 Nereis suecinea 80.4 72.3

Tellina agilis 5.6 1.0 Nephtys pieta 74.6 27.3

Nephtys pieta 5.2 1.2 Cancer irroratus 67.6 67.6

Jul 1983 5 grabs Tellina agilis 160.6 27.0 Nassarius triviUatus 903.4 463.3

Capitella spp. 34.8 7.7 Ceriantheopsis americanus 678.0 260.7

ephtyidae spp. 16.4 7.5 Diopatra euprea 333.4 204.3

Spiophanes bombyx 13.2 3.1 Tellina agilis 272.8 72.1

Ensis direetus 11.2 4.1 Nephtys pieta 164.0 56.3

Nephtys pieta 8.4 3.4 Rhynchocoela spp. 134.2 82.0

Tharyx acutus 8.2 3.5 Spiophanes bombyx 54.2 17.6

Ceriantheopsis americanus 7.0 1.6 Glyeera dibranehiata 29.0 17.7

Spisula solidissima 6.8 2.4 Ensis directus 22.4 8.1

Cerastoderma pinnulaturn 3.8 1.2 Cancer irroratus 21.2 10.3

Nov 1983 5 grabs Nephtys pieta 83.4 29.9 Diopatra euprea 832.2 482.1

Spiophanes bombyx 65.0 19.3 Ceriantheopsis americanus 603.4 294.3

Capitella spp. 33.2 23.3 Nassarius triviUatus 495.0 303.6

Eehinaraehnius parma 22.8 10.2 Nephtys pieta 112.2 62.0

Ampelisea vadorum 18.4 13.1 Spiophanes bombyx 102.6 36.9

Parougia caeca 14.4 4.7 Asabellides oculata 47.4 20.1

Nephtyidae spp. 11.6 11.6 Capitella spp. 19.8 12.7

Asabellides oculata 11.2 4.1 Rhynchocoela spp. 16.0 15.0

Nereis suecinea 10.6 3.9 Nereis suecinea 13.2 8.8

Thm'Yx spp. 8.4 2.2 Tellina agilis 12.6 3.4

Aug 1984 5 grabs Spiophanes bornbyx 110.8 45.5 Nassarius triviUatus 604.4 453.6

Tellina agilis 34.8 10.4 Tellina agilis 391.2 145.2

Nephtys pieta 24.4 7.6 Spiophanes bombyx 242.2 135.7

Capitella spp. 21.6 20.1 Capitella spp. 181.6 179.9

Tharyx acutus 8.4 4.6 Nephtys pieta 119.2 52.3

Exogone hebes 3.4 1.7 Ceriantheopsis amerieanus 79.8 48.0

Ceriantheopsis amerieanus 1.8 0.6 Cancer i"roratus 54.8 24.7

Nassarius tnviUatus 1.6 0.9 Ensis direetus 19.8 10.5

Ensis di"eclus 1.6 0.7 Pherttsa affinis 15.4 9.4

Cancer i"roratus 1.6 0.5 Diopatra cuprea 9.2 9.2
20 NOAA Technical Report NMFS 103

Table 6 (continued)

No. of Density Biomass (mg)

Dale samples Species Mean E Species Mean SE

Jul1985 5 "rabs SpiophaneJ bombyr 111.4 46.1 Spiophanes bombyx 1130.0 591.6

Telbna agilis 51.0 10.2 Diopatra L"ltpT~a 545.4 458.3

Nephtyidae spp. 29.4 7.5 JVwsatius trivitlatus 333.0 172.8

Unciola inomta 23.2 9.6 Nrphtys picta 290.6 65.8

JVephtys pittJ. 21.4 5.2 CeriantheopslS ameriwnus 249.6 115.7

Tharyxspp. 14.8 5.5 Tellina agilis 174.2 60.5

Edutea ttitoba 12.8 2.3 Lumbrinl'Tls aClcularwT. 130.4 49.6

ranar irroralus 9.6 3.4 Cancer it'romlus 85.2 31.2

Capiiella spp. 8.2 2.2 Edotca trilvlla 63.0 14.4

Tharyx aeu tus 7.8 5.2 Unao/a irmr'lla 49.8 24.8

Oct 1985 2 ~rabs Asabcllidps nrulata 71.0 18.0 ('VGUrLTlUS tri1JiUatus 2262.5 1537.5

Tharyx spp. 10.0 7.0 Diopatm wprw !\78.0 235.0

Nephtys picta 9.0 7.0 Pherti..m aJfinis 295.0 294.0

Nassarius trivittatus 7.0 5.0 Cen1tntheopsiJ arne,icanus 127.0 127.0

Pherusa aJfinzs 6.5 1.5 Cancer irroratus 103.0 103.0

Spiophanes bumbyx 6.0 0.0 Lamilrincns aawlar'um 52.5 52.5

Nereis mccintu 5.0 5.0 Nephtys pieta 48.0 :15.0

Mysplla planuta 5.0 5.0 Rhynchocoela spp. 40.0 25.0

Exogone hebes 3.0 3.0 Asabcllides owlata 14.0 7.0

Diopatm cuprea 3.0 0.0 Nereis SULClnliu 8.5 8.5
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 21

--------- ------ -,

Table 7

Mean densities and biomasses per 0.1 m with standard errors and numbers of replicate samples analyzed, for l.Le

numerically dominant species at Station 4 on each sampling date.

No. of Density Eiomass (mg)

Dale samples Species Meall SE Species Mean SE

Dec 1979 5 grabs Amastigos caperatus 772.2 380.8 ThaTyx aeutus 215.2 132.7

ThaTyx aeutus 452.6 284.7 Amastigos caperatus 203.6 13'\. I

Spisula solidissima 67.1 10.!! Nephtys picta 169.4 51.3

Nephtyidae spp. 43.6 69 Echinarachnius paTma 145.4 143.7

Tellina agilis 41.2 12.8 Tellina agilis 79.6 35.5

Phomnis m'chitecta 33.€ 21.7 Glycera dibranchiata 43.2 10.F.

Spiophanes bombyx 2~.6 6.1 Spisula solidissima 38.4 8.6

Nephtys picta 26.8 8.1 Lumbrineris acieulamm 32.0 13.8

Glycera spp. Uuv.) ]3.0 22 Rhynchocoela spp. 26.6 7.0

Echinarachnius panna It.:! 5.9 Magelona riojai 26.0 21.1

Jul1980 4 grabs ThaTyx awtus 776.0 291.5 Cancer borealis 22750.0 22750.0

Nueula proxima 295.0 272.7 Nucula pmxima 1635.8 1548.0

Tellina agili 5 284.~ 94.6 Tellina agilis 1]]8.5 408.9

PolyduTa ligm 242.3 161.1 Nephtys incisa 664.5 445.7

Amastigos capemtu> 159.3 lOll Phorvms a,-chitecta 428.8 427.1

Phyllodoce (anatides) mucosa J44.5 86.0 CanceT inomtus 412.3 266.0

Phomnis architecta 71.8 69.4 Thm yx aeutus 336.3 165.1

Mediomastus ambiseta ~6.3 46.3 Lumbrineris acieularum 238.8 88.6

Spiophanes bombyx 27.5 14.5 Rhynchocoela spp. 223.5 128.9

Rhynchocoela spp. 27.3 ll.S Glycera dibranchiata 178.8 ll8.0

Dec 1980 5 grabs Nephlyidae spp. 34.4 ]2.0 Ensis directus 506.4 503.4

Tellina agilis 26.4 5.6 Lunatia heros 258.0 258.0

Spiophanes bombyx 24.2 9.8 Tellina agilis 108.2 30.6

Tanaissus liljeborgi 20.8 12.4 Nephtys bucera 100.4 27.5

Nephtys bucera 14.6 4.0 Sthenelais limicola 64.4 57.4

Rhepoxynius hudsoni 10.2 4.9 Rhynchocoela spp. 32.8 22.7

F.nsis diTectus 7.4 1.2 Cmngon septemspinosa 25.8 24.1

Pseudunciola obliquua 7.4 5.3 Nephtys picta 25.4 12.2

Magelona riojai 7.2 3.7 Spisula solidissima 24.8 24.8

Rhynchocoela spp. 2.6 1.0 Lumbrineris acieularum 15.0 12.1

Aug 1981 5 grabs Nephtys picta 79.0 22.0 Spisula solidissima 986.6 849.9

Pseudunciola obliquua 60.8 33.0 Cancer irroratus 595.8 389.9

Spiophanes bombyx 48.0 14.0 Ovalipes ocellatus 568.2 212.9

Tharyx aeutus 41.8 40.6 Nephtys picta 205.2 68.7

Tellina agilis 30.6 7.4 Tellina agilis 180.2 38.3

Tanaissus liljeborgi 29.6 15.8 Ensis diTectus 164.0 101.2

Spisula solidissima 23.6 14.1 Lumbrineris acicttlarum 127.8 75.1

Phoxocephalus Itolbolli 16.8 15.6 Glycera robusta 100.8 89.4

Pamugia caeca 15.6 14.9 Spiophanes bombyx 89.8 24.2

Prionospio steenstrupi 10.4 10.4 Spio filicornis 51.4 51.4

Jan 1982 5 grabs Amastigos capl'Tatus 781.2 345.1 Ensis directus ]]94.8 614.3

Tellina agilis 600.4 220.3 Lumbrineris acieularum 521.4 350.0

Nephtyidae spp. 141.6 21.7 Tellina agilis 415.4 165.3

Mediomastus ambiseta 118.6 84.6 Pitar morrhuanus 153.8 32.2

Tharyx aeutus 110.8 73.9 Phoronis architecta 143.6 88.8

I Pitar mOTrhuanus 59.8 29.3 Mediomast1lS ambiseta ]]4.0 108.5

Spiophanes bombyx 40.4 7.3 Nephtyidae spp. 103.4 22.1 j Nueula proxima 31.0 14.5 Glycera dibranchiata 94.8 30.7

PhoTonis architecta 25.4 16.7 Pandora gattldiana 87.0 87.0

Sthenelais limicola 10.8 3.8 Sthenelais limicola 75.2 21.8
22 NOAA Technical Report NMFS 103

Table 7 (continued)

No. of Density Biomass (mg)

Date samples Species Mean SE Species Mean SE

Aug 1982 5 grabs Mytilus edulis 551.6 536.9 Mytilus edulis 1859.6 1560.4

Amastigos caperatus 97.4 60.8 Cancer irroratus 809.4 765.9

Tellina agilis 57.0 lU; Tellina agilis 345.4 104.8

Nephtys picta 38.6 9.8 Ophioglycera gigantea 324.0 324.0

Tharyx acutus 27.2 22.3 Pitar morrhuanus 276.4 275.7

Polydora socialil 18.6 17.6 Nephtys picta 242.0 19.2

Harmothoe extenuata 15.2 15.0 Asterias vulgaris 236.8 236.8

Parougia caeca 10.8 4.6 Lumbrineris acieularum 141.0 118.9

Spisula solidissima 10.6 3.6 Glycera robusta 81.4 79.2

Scalibregma injlatum 5.2 5.0 Glycera dibranchiata 78.4 29.9

Jul 1983 5 grabs Tellina agilis 53.0 16.3 Lumbrineris acieularum 244.4 72.8

Nephtyidae spp. 27.2 5.5 Tellina agilis 149.0 60.6

Goniadella gracilis 23.8 15.2 Nephtys picta 59.4 27.3

Chiridotea tuftsi 8.0 4.6 Rhynchocoela spp. 48.2 15.9

Nephtys picta 6.8 3.4 Glycera dibranchiata 38.8 21.6

Caulleriella cf killariensls 6.4 4.2 Nephtys bucera 35.8 11.7

Spisula solidissima 6.2 1.2 Yoldia spp. 34.4 34.4

Spiophanes bombyx 5.2 1.9 Cancer irroratus 31.4 15.2

Pleudoleptoeuma minor 4.4 1.6 Glycera robusta 22.6 22.6

Hemipodus roseus 4.2 4.2 Spisula solidissima 19.4 6.3

Aug 1984 5 grabs Spiophanes bombyx 26.0 7.9 Nephtys bucera 187.4 78.4

Nephtys picta 11.4 4.0 Lumbrineris aricularum 106.6 79.3

Goniadella gracilis 9.0 8.8 Spiophanes bombyx 77.2 34.1

Pseudunciola obliquua 8.2 8.0 Nephtys picta 73.0 42.7

Tellina agilis 7.4 1.3 Tellina agilis 66.6 26.7

Spisula solidissima 7.0 5.3 Nassarius triuittatus 55.8 39.2

Glycera spp. Uuv.) 6.2 l.l Ensis directus 39.8 24.5

Tanaissus liljeborgi 6.2 6.0 Cancer irroratus 26.0 26.0

Caulleriella cf killariensis 6.0 2.6 Glycera spp. Uuv.) 21.6 11.2

Tharyx aeut.us 3.0 1.5 Glycera dibranchiata 21.0 10.7

Jun 1985 5 grabs Pseuduncwla obliquua 85.2 58.3 Tellina agilis 197.4 84.3

Tellina agilis 42.4 14.5 Lumbrineris acieularum 156.2 79.6

Nephtyidae spp. 17.0 5.2 Glyrera dibranchiata 87.0 70.4

Tanaissus liljeborgi 14.0 3.5 Spisula solidisslma 46.8 15.4

Caulleriella if killariensis 9.8 2.7 Rhynchocoela spp. 30.4 12.0

Spisula solidissima 7.8 3.4 Nassarius triuittatus 29.0 29.0

Rhepoxynius hudsoni 6.6 2.8 Pseudunciola obliquua 27.6 14.5

Spiophanes bombyx 6.2 2.0 Nephtys picta 26.0 7.7

Ensis directus 5.4 1.7 Nephtyidae spp. 21.4 4.8

Magelona riojai 5.2 3.3 Spiophanes bombyx 15.4 6.1

Oct 1985 2 grabs Nephtys picta 245 11.5 Nephtys picta 212.0 39.0

Tellina agilis 24.5 3.5 'Tellina agilis 122.0 70.0

Spisula solidlssima 17.5 0.5 Spisula solidissima 96.5 71.5

Caulleriella cf killariensis 16.5 11.5 Arctica islandica 95.0 95.0

Tharyx aeutus 14.5 14.5 Lumbrineris acicularum 77.5 15.5

Magelona rosea 7.0 7.0 Glycera dibranchiata 67.0 52.0

Chiridotca tuftsi 4.5 2.5 Je.·nsis directus 55.0 55.0

Rhynchocoela spp. 4.0 U.O Nassarius trivittatus 46.0 46.0

Lumbrinms acieularum 4.0 3.0 Rhynchocoela spp. 15.0 7.0

Goniada maculata 3.0 2.0 Caulleriella cf killariensis 6.0 4.0
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 23

Table 8

Mean densities and biomasses per 0.1 m with standard errors alld numbers of replicate samples analyzed, for the

numerically dominant species at Slation 26 on each sampling date.

No. of Density Biomass (mg)

Dale samples Species Meal! SE Species Mean SE

Dec 1979 5 grabs F.chinamc!mius parma 56.6 19.2 Eehinamehnius panna 16846.0 5403.9

Spisula solidismna 10.2 4.7 CeriantheopslS americanus 331.8 212.5

Nephtys pieta 9.2 4.0 Lumlnineris aciwlarum 83.8 24.9

Lumblineris acicularum 7.4 1.2 Nephlys buum 58.2 43.3

Nephlyidae spp. 6.4 2.4 Spisula solidissima 40.6 23.4

Tellzna agilis 4.6 2.2 Nephtys picta 25.2 15.4

Caulle,.iella cf killu.nensis 4.2 1.1 Cmngon septemspinosa 24.0 12.8

Ceriantheopsis arnenWl£US 3.6 2.0 Rhynchocoela spp. 19.0 16.8

Pholis macrocoxa 3.6 1.0 Tellina agilis 14.2 6.8

Parougia caem 3.2 1.4 Astarte castanea 13.8 11.9

Jul 1980 5 grabs J;·,.hinarachr;ius parma 27.8 9.7 F.chinaraehnius panna 19332.1 7682.7

Nephtys plda 12.7 2.5 Ceriantheopsis americanus 1311.9 0.4

Crnia'ltizeopsis americanus 12.2 0.6 Spisula solidissima 432.3 427.0

l.umbri'1eris acieularum 8.2 0.6 AstaTte castanea 271.3 207.0

Cancer iTToralus 7.3 2.2 Lumbrineris acieularum 241.4 36.4

ThaTyx donobmnchialis 2.8 0.9 Nassarius trivitwtus 92.4 92.4

RhY'1chL'coela spp. 2.3 1.0 };uclymene zonalis 65.7 63.6

Tellina agilis 2.3 1.5 Cancer inomtus 51.7 16.8

t;uclymene zon(.</ls 2.0 0.8 Rhynchocoela spp. 49.5 29.3

Glyceru dibTancltiata 1.6 0.6 Clymenella tOTquata 48.0 48.0

Dec 1980 5 grabs EchmarachnlUs !'arma 36.3 11.5 t;chinarachnius parma 34208.9 14360.2

Gonzadelia gracilis 16.4 9.2 Ce1iantheopsis americanus 1845.1 626.4

Spisultl lO11dl,sima 12.4 8.7 Cancer inomtus 1229.1 1229.0

Ceriantheopsis americanus 7.6 2.0 AstaTte undata 486.9 486.8

Clycem spp. Uuv.) 7. \ 3.3 Astarte castanea 449.0 392.1

L71mbTme,is acicularzi1li 6.4 1.8 Lumbrineris aciwlarum 385.8 129.2

Aglaophamu5 igalis 6.1 1.7 Rhynchocoela spp. 81.4 73.7

Pelecypoda spp. (unidentifiable) 6.1 5.1 Nephtys picta fi6.1 22.6

Tharyx acut'~s 4.9 3.1 Nephtys bucera 64.4 64.4

Rhynchocoela ;,pp. 4.5 2.9 Sigalion arenicola 26.3 20.4

Aug 1981 5 grabs Coniadella gracilis 24.3 15.4 Echinarachnius parma 17019.8 13878.8

Hemipodus mseus 13.1 6.5 Ceriantheopsis americanus 821.3 373.9

Echinarachnius parma 10.3 7.2 Nassll1ius trivittatus 384.3 186.7

Lumbrine.is acicularum 10.0 4.5 Lumbrineris acicularum 254.0 27.9

Tharyx donobTanchialis 9.3 3.1 Nephtys bucera 162.4 96.9

Harmothoc extenuala 8.3 2.4 Lunatia l1iseriata 93. 1 93.0

Spiophanes boml7yx 7.9 3.8 Astarte caslanea 75.4 69.4

Ceriantheopsis ammcanus 3.3 1.3 Nephtys picla 59.7 18.1

Nephtys piela 3.3 1.0 Aglaophamus circinata 53.8 26.8

Nephtyidae spp 3.0 1.0 Cancer irromtus 33.7 14.9

Jan 1982 5 grabs Tharyx acutw 27.8 10.5 Echinamchnius parma 55693.4 10755 I

Hemipadus raspus 27.4 14.3 Spisula solidissima 29603.4 29600.4

Echinarachnms parma 24.6 4.1 Cenantheopsis americanus 1613.2 387.8

Rhynchocoela spp. 15.0 5.8 Astarte castanea 442.0 326.0

Coniadella gra~ilis 13.2 5.6 Nassarius l1ivittatus 221.0 221.0

Nephtyidae spp. 9.4 2.5 Rhynchocoela spp. 138.0 103.1

Spisula solidissima 8.4 2.8 Lumbrineris aeicularum 74.0 45.6

CeriantheC'psis americanus 5.6 1.5 Nephtys bucera 59.2 21.8

Parougia caeca 3.0 0.8 Nephtys picta 48.8 35.2

Nephtys bucera 2.6 0.7 Sigalion arenicola 29.6 21.8
24 NOAA Technical Report NMFS 103

Table 8 (continued)

'0. of Density Biomass (mg)

Date samples Species Mean SE Species Mean SE

Aug 1982 :; "Tabs Eehinaraehnius parma 71.9 13.0 Eehinaraehnius parma 12900.1 30175.3 " Tharyx acutltS 20.6 2.0 Spisula solidissima 9128.8 9117.8

ephtyidae spp. 10.1 4.4 Ceriantheopsis amerieanus 1079.4 502.6

Hemipodus roseus 9.7 9.1 Nassarius trivit/atus 383.3 157.9

Nephtys pieta 9.3 2.1 Cerastoderma pinnulatum 173.6 173.6

Tharyx dorsobranehialis 8.4 2.8 Nephtys pieta 86.4 20.1

Rhynchocoela spp. 8.3 4.4 Lumlnineris aeicularum 57.0 25.7

Goniadella gracilis 8.3 5.1 Rhynchocoela spp. 48.3 31.2

Harmothoe extenuata 5.5 2.7 Cancer irroratus 21.6 9.0

Ceriantheopsis americanus 5.3 1.7 Nephtyidae spp. 19.0 8.3

Jul1983 5 grabs Thm'Yx acutus 63.8 33.2 Eehinaraehnius parma 40284.2 15300.4

Goniadella gracilis 56.8 26.0 Ceriantheopsis amerieanus 1628.0 315.5

ephtyidae spp. 28.8 6.6 Astarte castanea 453.0 232.1

Eehinaraehnius parma 18.6 8.0 Lummneris acicularum 383.6 61.6

Tharyx dorsoaranchialis 15.4 5.9 Rhynchocoela spp. 74.0 57.8

Ceriantheopsis amerieanus 9.6 2.9 Tharyx marioni 41.6 41.6

Astarte castanea 9.6 4.1 Sigalion arenicola 30.0 27.1

Leitoscoloplos acutus 8.4 6.2 Nephtys lJueera 24.2 21.8

Lummneris acicularum 6.8 1.6 Tharyx acutus 19.8 13.3

Hemipodus roseus 3.4 3.2 Nephtys pieta 11.2 2.5

Aug 1984 2 grabs Cerionlhlopsis fl1T'e,icanus 34.0 24.0 Eehinaraehnius parma 33717.5 20079.5

Thar.yx acutu.\ 33.5 18.5 Ceriantheopsis amerieanus 1112.0 675.0

Goniadella :;rucihs 15.5 9.5 Nassarius trivit/atus 323.0 323.0

Echinaraeh'lliu 5 parma 14.5 8.5 Spisula solidissima 312.0 312.0

Caulleriella cf. killarimsis 7.0 6.0 Cimformia grandis 219.0 219.0

Spiophanes a01flby", 3.5 2.5 Nephtys IJucera 123.5 123.5

Tharyx dorsobranehialis 3.5 1.5 Lumarineris acicularum 123.5 123.5

ephtyidae spp. 1.5 0.5 Tharyx acutus 27.5 19.5

Glycera spp. Uuv.) 1.5 0.5 Astarte castanea 24.0 24.0

Ensis directus 1.5 0.5 Cancer irroratus 22.0 22.0

Jun 1985 5 grabs Echinaraehnius parma 42.6 5.5 Eehinaraehnius parma 10487.1 13261.5

Caulleriella tf killariensis 12.2 4.7 Ensis directus 595.6 589.4

Tharyx aeulu S 11.6 4.3 Ceriantheopsis americanus 503.0 201.8

Nephtys piela 7.4 3.2 Lummneris acicularum 476.4 53.1

Canter irroratus 6.2 0.9 Nephtys IJucera 174.6 130.3

Ncphtyidae spp. 5.6 3.5 Nephtys picta 85.2 35.6

Lumarineris n.ciwlarum 5.0 0.8 Cancer irroratus 45.8 11.6

Ceriantheopsis americanus 4.8 0.8 Astarte castanea 36.6 21.6

Ensis directus 3.8 l.l Rhynchocoela spp. 27.0 12.1

Tharyx dOTJobranehialu 3.0 1.0 Periploma leanum 21.0 21.0

Oct 1985 2 grabs Tellina agilis 43.0 30.0 Echinaraehnius parma 36130.0 31990.0

EchinaraehniusjJarma 30.0 2.0 Lummneris acicularum 122.5 121.5

Caulleriella cf killa'ritl'lsis 24.0 18.0 Ceriantheopsis americanus 75.5 10.5

Tharyxspp. 16.0 8.0 Cancer irroratus 36.0 36.0

Tharyx dursaaranehiahs 10.5 9.5 Astarte castanea 12.5 12.5

Magelana raseo 4.5 4.5 Tellina agilis 8.0 6.0

Spiophanes bombyx 4.0 4.0 Nephlys lJueera 7.0 2.0

ephtyidae spp. 3.0 3.0 Caulleriella cf killariensis 4.5 3.5

Nephtys aucera 2.0 0.0 Nephtys pieta 4.0 4.0

Syllides sp. I 1.5 1.5 Tharyxspp. 3.0 2.0
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 25

Table 9

l Mean densities and biomasses per 0.1 m with standard errors and numbers of replica~e samples analyzed, for the

numerically dominant species at Station 31 on each sampling date. --_._-- I

No. of Density Bioma~~ (mg) ! Date samples Species Mean SE Species Me:>n SE I ._------ ----------- Dec 1<)79 4 grabs Echinarachnius pa,-on" 552.8 106.3 Echinarachnius parma 'HI50.0 8350.4

PseudunC'tola obliquua 197.5 46.5 Lumbrineris aciwlarum 454_5 441.9

7(mailsu'liljeborK: 96.8 40.1 Acanthohaustorius spinOS1t:i 128_3 60_9

Byblis ,trrata 79.0 58.2 Byblis serrata 112.fi 78_:;

Telli~/.O agilis 40.5 21.8 Pseudunciola obliquua 72.8 16.2

Rhe-/Joxynius hud';(jni 38.8 17.9 Rhepoxynius hudsoni 61.0 n8

Tlwryx acu/us 29.0 2.7 Aglaophamus circinata 37.5 '\5.8

Exogone hebes 22.8 6.3 Pandora gouldiana 31.5 31.5

Aricidea (acesta) catherinae 22.3 5.5 Euclymene zonalis 27.8 ',_8

Corophium crassicorne 21.0 13.1 Nephtys incisa 21.5 ~1.5

.lui 191>0 5 grat>s Pseudunciola obliquua 135_3 45.4 Echinarachnius panna 2546.6 102'i9

7I:maissus liljeborgi 88.9 17.3 Ensis directus 284.Y 283.8

Exogone hebes 69.3 13.3 Byblis serrata ~41U 72.1

Byblis serrata 40.5 10.7 Nassarius trivittatus III I 111.0

Aoridac spp. Uu\,.) 29.3 29.2 Sigalion arenicola 76.4 32.7

Unciola ir-roratn 23.0 4.8 Unciola irrorata ~9.0 10.5

ATicidec (acesta) catherinae 20.6 4.6 Cancer irroratus 33.8 8.4

Coniadella gracilis 18.4 9.0 Rhepoxynius hudsoni 21.3 12.9

Caulle/irlln if. killuriensis 17.6 7.2 Drilonereis magna 18.9 13.8

Rhepoyynius Imdsoni 16.6 5.8 Pseudunciola obliquua 18.<) 2.8

Dec 1980 5 grabs Byblis senata 111.3 25.7 Echinarachnius parma 12436_0 6483.8

t;xogone hebes 37.0 10.3 Astarte undata 2:;80.1 2580.0

Tanaissus liljeboTgi 36.0 ll.O Nassarius trivittatus ~31.6 231.6

Pseudunciola obliquua 32.6 12.9 Byblis serrata 1!J1.6 63.5

Rhepoxynius hudsoni 29.0 2.7 Rhynchocoela spp. 145.9 140.6

Nephtyidae spp. 18.8 5.4 Ensis directus 130.4 129.4

Spiophane.' bombyx 17.3 2.8 Cirolana polita 63.8 35.7

Unciola il"Tomta 13.4 3.5 Cancer irroratus 60.0 36.8

Aricidea (acesta) catherina,' 13.3 4.2 Lumbrineris acicularum 46.8 24.5

Corophium cmssicorne 10.9 3.0 Euclymene zonalis 46.7 16.1

Aug 1981 5 grabs Pseudunciola obliiJuua 534.3 218.8 Echinarachnius pa'rma 2973.6 J87'7.5

Tanaissus izljeborg:. 82.9 20.7 Ascidiacean spp. 819.5 452.4

Comphium cmssicorne 68.1 13.6 Ceriantheopsis americanus 645.8 2 J 1.1

Exogone hebes 53.3 19.1 Sigalion aTenicola 156.1 92.2

F:chinamc"nius parmc 45.3 9.4 Corophium crasSlconu 153.6 111.8

RhP/Jo ynius hudsuni 31.0 100 Pherusa aJfinis 145.6 145.6

Aricidea (acesta) Cfltherinat: 27.3 8.5 Unicola inoratn 145.1 89.8

Euciymene zonalis 20.3 10.2 Spiophanlis bumbyx 84.4 20.6

Cauleriellu If killariensis 15.4 3.0 Psuedunciola obliquua 77.7 36.3

Ultiiola irrorata 14.7 3.] Rhynchocoela spp. 69.7 31.8

Jan 1982 5 grabs Tanmssus liljeborgi 310.0 67.9 F:(hinanz(hnius panna 20801.6 7236.3

Echinarachnius panna 185.8 32.1 Astarte mstenea 2385.0 2383.8

Pseudunciula obliquua 161.8 52.8 Ceriantheopsis americanus 1043.4 597.3

Exogene hebes 78.2 15.6 Cancer irromtus 129.4 79.5

Aricidea (acesta) cathClinae 36.4 7.4 Rhynchocoela spp. 120.6 66.5

CaullCliella c! killariensis 19.8 4.2 PseuduT'ciola obliquua 76.2 26.2

Tharyx acutus 14.8 8.9 Cirolana polita 47.0 47.0

Sliiophanes bombyx 11.2 2.7 Sthenelais limicola 36.8 14.7

Tellina agilis 10.8 4.3 Nereidae spp. 36.4 36.4

Polydora caulleryi 9.8 3.1 Tharyx acutus 24.8 13.7
26 ,NOAA Technical Report NMFS 103

Table 9 (continued)

No, of Density Biomass (mg)

Date samples Species Mean SE Species Mean SE

Sep 1982 5 grabs Pseudunciola obliquua 390.8 256.4 Echinarachnius parma 14277.3 5865.1

Tanaissus liljeborgi 70.5 32.8 Phoronius architecta 742.7 740.4

Echinarachnius pm'ma 65.1 25.6 Ceriantheopsis americanus 6I6.5 17.2

Phomnis architecta 62.0 60.3 Nassarius trivittatus 383.3 237.8

Euciymene zonalis 38.3 18.3 Lu natia triseriata 225.6 195.2

Exogene hebes 29.4 9.4 Rhepoxynius hudsoni 93.7 15.7

Clymenella torquata 28.6 18.2 Pseudunciola obliquua 91.4 57.2

Rhepoxynius hudsoni 26.3 5.1 Cancer irroratus 90.1 47.7

Corophium crassicorne 25.6 3.9 Clymenella t01'quata 66.1 41.6

Caulleriella cf killariensis 23.8 12.5 Pherusa a/finis 50.5 40.9

Jul 1983 5 grabs Pseudunciola obliquua 455.4 63.1 Echinarachnius parma %167.2 21809.7

Tanaissus liljeborgi 234.8 40.8 Lumbrineris acicularum 178.4 ll9.5

Exogene hebes 87.2 42.3 Ascidiacean spp. 160.8 160.8

Caulleriella cf killariensis 27.4 2.5 Nassarius trivittatus 83.0 83.0

Polydora caulleryi 23.6 17.8 Sigalion arenicola 81.4 18.8

Goniadella gracilis 19.6 6.6 Pseudunciola obliquua 64.2 4.2

Aricidea (acesta) catherinae 19.4 4.0 Rhynchocoela spp. 51.0 36.7

Echinarachnius parma 14.4 6.3 Cancer irroratus 39.8 14.4

Rhynchocoela spp. 9.2 3.3 Orbinia slllani 31.4 31.4

Rhepoxynius hudsoni 8.6 0.8 Aglaophamus circinata 23.0 17.6

Aug 1984 5 grabs Spiophanes bombyx 145.8 64.5 Echinarachnius parma 11087.6 7056.0

Pseudunciola obliquua 133.4 28.1 Ascidiacean spp. 1321.6 849.4

Exogone hebes 52.0 15.8 Ceriantheopsis americanus 670.4 292.3

Caulleriella cf killmiensis 34.8 10.3 Lumbrine1is acicularum 523.4 225.9

Tharyx acutus 24.4 6.1 Pherusa a/finis 185.2 129.4

Tanaissus liljeborgi 22.4 9.3 Spiophanes bombyx 157.6 87.3

Ceriantheopsis americanus 21.0 8.7 Pseudunciola obliquua 138.4 87.4

Ascidiacean spp. 15.4 8.2 Sigalion arenicola 107.6 39.6

Arcidea (acesta) catherinae 14.8 3.0 Tellina agilis 100.0 17.2

Phoxocephalus holbolli 13.8 7.6 Glycera dibranchiata 94.2 76.3

Jun 1985 5 grabs Nephtyidae spp. 26.4 3.1 Echinarachnius parma 13566.8 4145.4

Tellina agilis 25.8 5.8 PitaI' m01'Thuanus 2733.2 2727.7

Tharyx acutus 24.8 8.5 Nassarius trivittatus 473.6 193.4

Phoxocephalus holbolli 18.2 8.2 Ceriantheopsis americanus 306.2 104.5

Spiophanes bombyx 15.4 6.6 Tellina agilis 205.8 65.9

Harmothoe extenuata 10.8 4.1 Sthenelais limicola 88.8 38.0

Tharyx dorsobranchialis 10.4 6.7 Rhynchocoela spp. 83.4 22.3

Caulle1iella cf killariensis 10.0 5.7 Spiophanes bombyx 48.6 31.4

Exogone hebes 9.4 5.1 Lyonsia hyalina 42.2 17.2

Ensis directus 8.0 1.7 Nucula proxima 41.2 16.2

Oct 1985 2 grabs Clyce1'a spp. 16.5 4.5 Echinarachnius parma 16950.5 16949.5

Tellina agilis 13.5 5.5 Nassarius trivittatus 212.0 212.G

Echinarachnius parma 6.5 2.5 Tellina agilis 38.0 35.0

Nuwla pruxima 5.5 5.5 Cancer irroratus 37.0 37.0

Asabellidcs oculata 5.0 2.0 Sigalion arenicola 21.5 21.5

Pseudunciola obliquua 5.0 1.0 Solen viridis 20.5 20.5

Aricidea (acesta) catherinae 4.0 1.0 Nucula proxima 18.0 18.0

Tanaissus liljeborgi 3.5 2.5 Pherusa a/finis 12.0 11.0

Exogone hebes 3.0 2.0 Rhynchocoela spp. ll.5 3.5

Poly cirrus eximius 3.0 0.0 Sthenelais limicola ll.O 11.0
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 27

Table 10

Mean densities and biomasses per 0.1 m with standard errors and numbers of replicate samples analyzed, for the

numerically dominant spe,:ies at Station 15 on each sampling date.

No. of Density Biomass (mg)

Date ;,amples Species Mean SE Species Mean SE

Dec 1979 5 grabs Ampelisca agassizi 1297.4 303.1 PitaT morrhuanus 31682 3158.0

Tharyx aetttus 881.6 271.5 Ampelisca agassizi 1914.4 633.9

Euchone incolor 571.6 274.0 Nephtys incisa 1085.6 275.2

Cossura longocirrata 360.8 71.6 Asterias spp. a 384.0 384.0

Levinsenia gracilis 248.4 38.2 Chone iTlfttndibuliformis 355.2 135.0

Euclymene zonalis 172.2 40.8 Nicolea venustula 341.6 121.4

Aricidea (acesta) catherinae 156.6 58.8 Periploma papYTatium 315.0 175.3

Mediomastus ambiseta 151.2 58.0 Tharyx aeutus 292.4 67.3

Prionospio steenstrupi 145.2 26.9 Rhynchocoela spp. 276.6 96.1

Hucula delphinodonta 113.4 51.0 Nucula delphinodonta 273.8 192.4

Jul1980 5 grabs Ampelisca agassizi 1475.5 235.0 Ampelisca agassiz; 2811.3 404.8

Euchone elegans 740.6 717.1 Nephtys incisa 1166.8 448.8

Tha1}x dorsobranchialis 583.3 347.2 Thyasira trisinuata 482.3 258.8

Photis macrocoxa 231.4 130.0 Solemya borealis 472.0 472.0

Mediomastus ambiseta 170.4 80.0 Chone infundibuliformis 392.9 273.5

Nucula delphinodonta 161.9 49.9 Pherusa affinis 381.3 60.5

Levinsenia gracilis 154.3 79.7 Leptocheirus pinguis 314.5 121.3

Lumbrineris hebes 132.2 52.2 Anobothrus gracilis 272.9 138.2

Euchone incolor 119.7 61.3 Tharyx dorsobranchialis 242.4 145.8

Diastylis abbreviata 100.5 49.5 Mediomastus ambiseta 238.3 114.9

Dec 1980 5 grabs Ampelisca spp. 235.6 58.5 Astarte undata 2429.3 2172.0

Ampelisca agassiii 145.4 28.1 Astarte crenata subequilatera 2244.1 2244.0

Leptocheirus pinguis 129.4 54.5 Nephtys incisa 779.8 486.6

Nucula delphinodonta 115.1 32.5 Leptocheirus pinguis 562.5 180.4

Ninoe nigripes 49.6 13.6 Chone infundibuliformis 474.5 99.5

Erichthonius rubricornis 41.0 17.3 Cyclocardia borealis 456.3 145.9

Unciola irrorata 37.8 9.2 Cancer irroratus 436.9 285.2

Scalibregma inflatum 37.3 6.7 Scalibregma inflatum 394.6 73.1

Lumbrineris hebes 35.4 12.7 Ninoe nigripes 361.6 164.8

Nucula proxima 32.0 5.2 Periploma papyratium 352.6 83.6

Aug 1981 5 grabs Euchone incolor 1245.5 262. ) Nephtys incisa 1985.1 320.3

Tharyx dorsobranchialis 437.6 123.4 Thyasira trisinuata 1795.8 331.9

Nucula delphinodonta 380.3 128.0 Pherusa affinis 1130.8 456.3

Ampelisca agassizi 283.0 161.0 Yoldia sapotilia 1101.4 345.7

Cossura longocirrata 252.1 97.6 Nueula delphinodonta 809.5 311.5

Lumbrineris hebes 212.0 60.3 Pentamera calcigera 706.7 432.0

Photis macrocoxa 180.1 40.5 Nucula proxima 543.3 286.8

Levinsenia gracilis 174.4 47.3 Edotea triloba 390.6 62.0

Aricidea (Acesta) catherinae 160.3 52.7 Periploma papyratium 372.6 164.1

Polydora socialis 147.4 44.5 Ceriantheopsis americanus 342.3 69.8

Jan 1982 5 grabs Ampelisca agassizi 1247.8 189.2 Pitar morrhuanus 5654.4 3467.0

Euchone incolor 861.8 152.1 Ampelisca agassizi 2264.4 332.8

Lumbrineris hebes 169.0 17.7 Nephtys incisa 1660.0 589.4

Tharyx dorsobranchialis 159.2 29.7 Axius serratus 1370.2 1370.2

Ninoe nigripes 136.4 7.7 Asterias vulgaris 986.8 986.8

Nucula delphinodonta 124.0 9.2 Glycera robusta 810.0 810.0

Maldanidae spp. 4 70.2 28.4 Ninoe nigripes 351.4 66.6

Levinsenia gracilis 60.8 9.0 Pherusa affinis 305.2 160.7

Photis dentata 59.6 13.4 Nicolea venustula 291.0 131.0

Erichthonius rubricornis 56.4 11.1 Periploma papyratium 177.6 42.1
28 NOAA Technical Report NMFS 103

Table 10 (continued)

o.of Density Biomass (mg)

Dale samples Species Mean SE Species Mean SE

Sep 1982 5 grabs Ampelisca agassizi 490.8 160.6 Nephtys incisa 1421.1 664.6

Euchone incalor 433.4 196.1 Ampelisca agassizi 710.5 253.4

Tharyx dorsobranchialis 292.3 ] 14.7 Periploma papyratium 404.0 114.4

Nucula delphinodonta 219.1 110.6 Ninoe nigripes 372.5 101.2

Polydora socialis 186.1 123.4 Yoldia sapotilla 353.5 226.0

Lumbrineris hebes 173.0 40.2 Nucula delphinodonta 311.5 168.0

Ninoe nigripes 136:7 16.5 Rhodine gracilior 2]8.4 117.6

Cossura longocirrata 99.] 29.3 Nucula proxima 209.3 131.7

Levinsenia gracilis 96.3 28.3 Chone infundibuliformis 162.4 76.6

Aricidea (Acesta) catherinae 75.7 32.] Thyasira trisinuata 142.] 126.5

Aug 1984 5 grabs Euchone incalor 420.0 192.7 Nephtys incisa 3069.6 618.1

A mpelisca agassizi 179.4 119.7 Rhodine gracilior ]388.8 442.3

Tharyx dorsobranchialis 152.4 77.3 Havelochia scabra 1281.8 1281.8

Prionospio steenstTZtpi 147.4 34.4 Glycera robusta ]045.0 1045.0

Aricidea (Acesta) catherinae 123.2 99.2 Ampelisca agassizi 502.6 229.5

Cossura longoci1'rata 109.2 36.4 Pitar morrhuanus 362.6 338.6

Nucula delphinodonta 107.4 64.9 Periploma papyratium 273.4 109.1

Lumbrineris hebes 99.4 35.7 Thyasira trisinuata 254.2 64.1

Levinsenia gracilis 89.8 45.1 Yoldia sapotilla 232.0 170.3

Ninoe nigripes 57.8 10.9 Driloneuis longa ]98.6 35.0

JUIl ] 9 5 5 grabs Ampelisca agassizi 392.4 73.] Arctica islandica 13533.8 13237.8

Eudorella pusilla ] 18.6 18.6 Ninoe nigripes 1332.8 230.7

Aricidea (Acesta) catherinae 57.8 16.7 Ampelisca agassizi 635.8 124.3

Ninoe nigripes 52.2 12.7 Chone infundibuliformis 460.2 321.7

Harpinia propinqua 49.4 12.5 Astarte undata 272.2 163.5

ephtyidae spp. 45.6 6.1 Nephtys incisa 201.8 85.9

Diastylis quadrispinosa 35.4 10.1 Pitar morrhuanus 198.8 105.0

Unciola irrorata 34.4 12.1 Periploma fragile 163.4 71.7

Crenella glandula 32.4 7.8 Cyclocardia borealis 151.0 58.9

Lumbrineris hebes 29.8 3.4 Diastylis quadrispinosa 146.4 44.2

Oct. 1985 2 grabs Ampelisca agassizi 358.5 30.5 Nephtys incisa 2743.0 1708.0

Nucula proxima 90.0 84.0 Ensis directus 606.0 606.0

Lumbrineris hebes 79.5 20.5 Ampelisca agassizi 579.0 170.U

Prionospio steenstrupi 74.5 22.5 Chone infundibuliformis 271.5 263.5

Levinsenia gracilis 62.0 22.0 Periploma papyratium 260.0 48.U

Tharyx dorsobranchialis 59.0 30.0 Pherusa affinis 239.0 91.0

Harpinia propinqua 57.0 20.0 Nucula proxima 179.5 171.5

Aricidea (Acesta) catherinae 37.5 26.5 Rhodine gracilior 111.5 61.5

Nucula delphinodonta 37.0 29.0 Lumbrineris hebes 108.5 19.5

Nephtys incisa 36.0 22.0 Thyasira trisinuata 87.5 25.5
Reid et aI.: Benthic Macrofauna of the New York Bight, 1979-89 29

Table 11

Mean densities and biomasses per 0.1 m with standard errors and numbers of replicate samples analyzed, for the

numerically dominant species at Station I on each sampling date.

No. of Density Biomass (mg)

Date samples Species Mean SE Species Mean SE

Jul 1980 1 grab Nueula proxima 2283.1 0.0 Antiea lslandica 25301.1 0.0

Phoronis arehi/eeta 842.0 0.0 Phoronis arehiteeta 5825.0 0.0

Thmyx aeutus 507.0 0.0 Nueula proxima 5617.0 0.0

Nephtys ineisa 160.\ 0.0 Ceriantheopsis amerieanus 3536.1 0.0

Photis macrocoxa 80.\ 0.0 Nephtys ineisa 3475.1 0.0

Cossura longoeirrala 67.1 0.0 Pherusa a/finis 2876.1 0.0

Teliina agilis 50.0 0.0 Tharyx aeutus 346.0 0.0

Mediomastus ambiseta 48.0 0.0 Ensis direetus 289.0 0.0

Prionospio steenstrupi 41.0 0.0 Teliina agilis 263.0 0.0

Ceriantheopsis americanus 37.0 0.0 Crangon septemspinosa 39.0 0.0

Aug 1981 1 grab Nucula proxima 1348.1 0.0 Nereis virens 9242.0 0.0

Tharyx aeutus 1112.1 0.0 Nueula proxima 7640.0 0.0

Phoronis arehiteeta 446.0 0.0 Nephtys incisa 5240.0 0.0

Cossura longocirrata 156.1 0.0 Phoronis arehiteeta 4020.0 0.0

Nephtys incisa 59.0 0.0 Ceriantheopsis amerieanus 3360.1 0.0

Cerianth eopsis americanus 48.0 0.0 Tharyx aeutus 680.0 0.0

Prionospio steenstrupi 40.0 0.0 Yoldia limatula 634.0 0.0

Mediomastus ambiseta 38.0 0.0 Pherusa a/finis 383.0 0.0

Pherusa a/finis 30.0 0.0 Cerastoderma pinnulatum 31.0 0.0

Lwnbrineris hebes 15.0 0.0 Phyliodoee (Anatides) mucosa ~().O 0.0

Aug 1982 2 grabs Tharyx aeutus 4492.5 502.5 Nueula proxima 15587.0 2716.0

Nueula proxima 4086.1 140.0 PhOTonis arehiteeta 7535.5 413.5

Cossura longocirrata 895.5 19.5 Nephtys incisa 4084.1 927.0

Phoronis arehiteeta 737.0 ll2.0 Cerianlheopsis amerieanus 3328.1 ~~4.G

Mediomastus ambiseta 364.0 52.0 Pherusa a/finis 1897.1 54.0

Euehone ineolor 174.6 82.5 Thmyx aeutus 932.5 256.5

Prionospio steens/rupi 124.1 10.0 Edwardsia elegans lR4.1 89.0

Nephtys incisa 73.1 4.0 Yoldia sapotilia 159.6 159.5

Lurnbrineris hebes 61.5 9.5 Mediomastus ambise/a 123.1 10.0

Ninoe n;f!'ipes 51.0 0.0 Lu:mbrioneris hebes 112.0 14.0

Sep 1983 2 ~rabs Nucuta proxima 4331.0 205.0 Nueula proxima 19675.5 184.5

ThaTyx aeutu. 2615.5 747.5 Antieu iStarid/.ca 122Y9.0 12299.0

Prionospic stews/rupi 1222.5 386.5 Phor"nis anhitato 11467.5 1932.5

Cossura tongocirrata 905.0 124.0 Pherus(}. a/finis 5442.5 1280.5

MedlOmas/us ambise/c. 460.5 1.5 Nrph/y~ znd,a 4520.5 205.5

PhOTOnzs archiieela 459.5 76.5 Ceriantheol'sis am~rieanus 3230.5 170.5

Tharyx dorsobranchialis ll7.5 ll.5 Tharyx aewU> 2021.5 568.0

Pherusa a/finis Y4.5 32.5 Rhynehoeoela 769.5 719.5

Cl!1iantheopsis american1l.' 88.0 5.0 Prionospio steenstrupi 368.5 145.5

Lumbrineris hebes 62.5 9.5 Yoldia tit/ltula 275.0 275.0

Aug 1984 2 grabs Nueula proxima 3650.5 1171.5 Nucula proxima 16432.5 5730.5

Tharyx aeutus 1330.5 43.5 Pherusa a/finzs 7866.0 2722.0

Cossura ton/{ocirrata 321.5 146.5 Phomnis arehitee/a 6584.5 1328.5

Phoronis anhitee/a 234.0 9.0 Nereis virens 5026.0 5026.0

Prionospio steenstmpi 197.0 21.0 Nephtys ineim 4434.0 652.0

Mediomas/tls ambiseta 78.0 17.0 Cerian/heopsis amerieanus 2877.0 652.0

r:eriantheopsis amerieanus 69.0 28.0 Glyeera robusta 990.0 990.0

Pherusa a/finis 59.:> 5.5 Tharyx aeutus 963.5 13.5

Nephtys incisa 55.5 7.5 Pitar mOTThuanu~ 798.5 784.5

Lumbrineris hebes 44.0 5.0 Goniada norvegiea 648.0 648.0
30 NOAA Technical Report NMFS 103

,._-------------------------------------------------,

I

I Table 11 (continued)

8388.5

351.0

82.0

35.0

672.0

8.0

176.0

0.5

58.5

28.5

12313.5

8122.0

2148.0

1429.0

1428.0

254.9

184.0

70.5

58.5

52.5

50.5

687.5

88.5

127.0

120.0

44.0

23.0

lO.O

16.5

6.0

2592.5

716.5

191.5

191.0

169.0

102.0

47.0

41.0

37.5

31.0

Nucula proxima

Tharyxspp.

Phoronis architecta

Tharyx acutus

Cossura longocirrata

Pherusa affinis

Mediomastus ammseta

Nephtys incisa

Lumbrineris hebes

Ceriantheopsis americanus

Date

Oct 1\-185 2 grabs

No. of Density Biomass (mg)

sample> Species Mean SE Species Mean SE ------ ._---_:---------------------~-----._--------

Pherusa affinis

Nucula proxima

Nephtys incisa

Ceriantheopsis americanus

Phoronis architecta

Rhynchoceola spp.

Tharyxspp.

Edwardsia elegans

Nassarius trivittatus

Tharyx acutus

Dominant Species

Lists of the ten most abundant species in both numbers and biomass for each station and sampling have

been prepared. These lists complement the above

treatments of higher taxa, such as amphipods, and

the broader analyses (species richness, clustering) by

revealing which species are responsible for any spatial-temporal trends detected. Analysis of dominant

species can indicate whether these trends are due to

pollution-sensitive or tolerant species. The dominant

species lists can be combined with information on

fish diets (summarized in Reid and Steimle 1988) to

estimate the forage value of a site's macrobenthos

and whether that value has been altered.

Dominant species for the six NEMP stations (4, 6,

7, 15, 26, and 31) (Fig. 1) are emphasized because

these stations have more samplings, including winter

collections, and greater replication (usually five

grabs). For Station 6, there is the added advantage of

having monthly data available for summer 1986 and

1987 and counts for the polychaetes, Capitella spp.,

for summer 1988. The NEMP stations provide fairly

good spatial coverage of the overall study area. They

also include most of the habitat types and faunal assemblages encountered. Data on dominant species

for the NEMP stations are given in Tables 5-10. Conspicuously absent from the NEMP siteS is an assemblage consistently found in and near the

Christiaensen Basin, with high densities and biomasses of polychaetes, bivalves, and an anthozoan

(Pearce et al. 1981; Steimle 1985). The high abundances in this assemblage may be due to organic enrichment (Boesch 1982). Lists of dominant species

from Station 1 (Table 11) illustrate trends in the "enriched" assemblage. Tables of dominant species for

other non-NEMP stations are available on request.

Some noteworthy features of Tables 5-11 are discussed below.

Station 6 (sewage sludge accumulation area-Table

5)-As noted above, this area apparently contained

the most altered assemblage of any station. The fauna

is characterized by Capitella Spp.l, which Awere the

numerical Gominants in eight of the 13 samplings at

Station 6, with a peak mean density of 5604 individuals/0.1 m2. Capitella spp. were rarely the most abundant taxa outside of Station 6, and maximum density

never exceeded 678/0.1 m2 elsewhere.

There may be a seasonality in abundance of

Capitella spp. at Station 6 (Fig. 10). In winter samplings from 1979 through 1983, Capitella spp. were

the top dominant only once (November 1983), and

mean densities ranged from 1-415/0.1 m2 • Conversely, Capitella spp. were the top dominants in all

summer samplings from 1980 through 1985, though

abundances were quite variable (x = 87-5604/0.1 m2).

Where there were multiple samplings in a given summer, shorter-term fluctuations are indicated. There

were large increases in abundance between July and

August 1983 and between July and September 1987,

I Capitella capitata is widely used as an indicator of organic enrichment, although it also reaches high densities in response to

other disturbances, such as defaunation (Caracciolo and

Steimle 1983). Capitella capitata is actually a complex of several

species which are morphologically similar but have distinct genomes and life histories (Grassle and Grassle 1976; Grassle et

al. 1987). The differences may be important in interpreting

population fluctuations and in using Capitella spp. as pollution

indicators. The Capitella specimens from our sampling that

have been examined closely appear to resemble species 1a in

Grassle and Grassle (1976) (Judith Grassle, Marine Biological

Laboratory, Woods Hole, MA 02543, pers. commum., November 1988). Species 1a produces a relatively large number (200-

2000) of small eggs which remain in the plankton for several

days, whereas the planktonic phase is shorter or absent in most

other Capitella species examined to date. Since most of our

specimens have not been positively identified, they will be referred to as Capitella spp.
Reid et aI.: Benthic Macrofauna of the New York Bight, 1979-89 31

and a major decrease between July and August 1986.

Densities were low (<10/0.1 m2) in July through September 1988.

One hypothesis to explain the sudden changes is

that populations of Capitella spp. are limited by deteriorating water quality (e.g., high temperature, low

dissolved oxygen, concentrations of hydrogen sulfide), for variable periods during at least some summers. Recurrent hypoxia, and occasional sulfide generation, have been observed at Station 6 (Andrew

Draxler, NMFS, Sandy Hook Laboratory, Highlands,

NJ 07732, pers. commun., November 1988). Some

species of the genus Capitella have been reported to

have only a moderate tolerance for hypoxia (Reish

1970). Tsutsumi (1987) documented the disappearance of a dense C. capitata population from an organically polluted cove in Japan in mid-summer; the

population did not begin to reestablish itself until

late October, when dissolved oxygen increased and

sediment sulfide content began to fall. The factors

controlling Capitella spp. densities at Station 6 will be

examined by comparing densities to physical/chemical data (e.g., temperature, oxygen, redox potential,

and sulfide) being collected in the sludge dumpsite

Recovery" study (Environmental Processes Division,

Northeast Fisheries Center 1988), and also to the limited historical data available.

There are of course other possible causes for the

observed variability, perhaps acting in concert with

changing water quality. Laboratory populations of

Capitella spp. are known to have oscillations in abundance, which have been linked to declining food supplies and overshooting a habitat's carrying capacity,

even under constant conditions (Chesney 1985).

Much of the variability is undoubtedly due to the

combination of spatial patchiness in densities of

Capitella spp. and inexact station relocation for multiple sampling, as shown by the wide confidence limits in Figure 10. In October 1985, no Capitella spp.

were found at Station 6. There were also no

rhynchocoels, normally a top dominant in biomass

and perhaps more useful than Capitella spp. in characterizing the sludge accumulation area, since

rhynchocoels generally did not show such large shortterm fluctuations (Table 5). The variability of the Station 6 assemblage makes it difficult to use in monitoring changes following phaseout of dumping.

Station 7 (sewage sludge dumpsite-Table 6)-As in

the sludge accumulation area, Capitella spp. were numerical dominants in all Station 7 samplings from December 1979 through June 1985 and then were not

found in the last sampling in October 1985. Densities

were less variable over time than those at Station 6,

and peak abundances were about an order of magnitude less. Similarly, rhynchocoels were generally a biomass dominant as at Station 6, but were considerably

less abundant at Station 7. The lower densities are

taken as an indication that the effects of organic enrichment in the dumpsite itself were fairly small; the

dumpsite is not a markedly depositional area, as indicated by its relative coarse, low-carbon sediments

(Table 1). Conspicuous among other biomass dominants were the anthozoan Cenantheopsis amencanus

and polychaete Diopatra cuprea, both large tube-dwelling species whose abundances may have been enhanced by the sludge input; both were less abundant

at Station 22, 5.6 km east of Station 6 and with similar

depth and sediments. There were no obvious faunal

trends indicative of changing effects of either organic

enrichment or toxicants over the study period.

Station 4 (New Jersey Inshore, 2.4 km NNE of

dredged material dumpsite - Table 7-The fauna was

variable, with no consistently dominant species. This

may be related to the sediment patchiness we have

encountered at Station 4, which in turn is probably

due in part to the history of past dumping. (We have

found no distinctly altered assemblage near this

dumpsite comparable to that of the sewage sludge

accumulation area.) Early collections at Station 4 had

high densities of several species, e.g., the polychaetes

Amastigos caperatus and Tharyx spp. In 1983-85 there

were no very abundant species, and the overall fauna

was sparse.

Station 26 (New Jersey Offshore, 24.8 km E of central

NJ coast-Table 8)-This station was situated at the

center of the 1976 hypoxia event off New Jersey

(Steimle and Radosh 1979). The benthos appeared to

have largely recovered by the beginning of this study

(unpubl. data). The fauna was numerically sparse

throughout the study period with no obvious trends

over time and no consistently abundant species, except perhaps for the sand dollar Echinarachnius

parma, which was always the overwhelming biomass

dominant. Steimle (1990) gives a more detailed analysis of biomass data for E. parma and other species at

the NEMP stations.

Station 31 (Long Island Offshore, 25.0 km SSE of

Fire Island Inlet-Table 9)-As a rule, this station was

dominated numerically by crustaceans, including the

amphipods Pseudunciola obliquua, Byblis serrata,
32 NOAA Technical Report NMFS 103

Corophium crassicorne and Rhepoxynius hudsoni, and

tanaidacean Tanaissus liljeborgi. Abundances of all

these species decreased between 1979 and 1985. The

sand dollar was always the biomass dominant, usually

by more than an order of magnitude, with levels

roughly comparable to those at Station 26 (above)

and wi th no clear trend between 1979 and 1985.

Station 15 (Outer Hudson Shelf Valley, 48.7 kIn SE of

Sewage Sludge Dumpsite-Tab1e 10)-This station had

been considered to represent the innermost part of

the Shelf Valley in which contaminant concentrations

and the macrofaunal assemblage were not influenced

by pollution (Boesch 1982). The significant decrease

in overall numbers of amphipods at this station over

the study period was noted above. One species,

Ampelisca agassizi, was responsible for most of the decrease. A. agassizi was the numerical dominant in

seven of the nine samplings. Mean densities of over

1000 individuals/O.l m2 were found in several of the

early samplings, but all counts from September 1982

on were';;; 490/0.1 m2, with an apparent continuing

decline through 1985. The species is thought to be

useful for trend monitoring owing to its sensitivity

and natural stability of populations (Boesch et al.

1977; Radosh et al. 1978; Schaffner and Boesch

1982). The trend in A. agassizi is another indication,

along with the general decreases in numbers of total

species and amphipods discussed above, that some

natural or pollution-related changes have recently affected at least the sensitive components of the

benthos. Station 15 also had marked declines in several other species, e.g., the polychaetes Levinsenia

gracilis and Euchone incolor, and the bivalve Nucula

delphinodonta. Overall numerical abundance decreased here, as part of a trend that was widespread

but not universal among Bight stations (e.g., also apparent at 4 and 31 but not at 6, 7, or 26, or in the

"enriched" assemblage discussed next).

Station 1 (Northern Christiaensen Basin, 5.6 km NW

of Sewage Sludge Dumpsite-Tab1e 11 )-This station

had an "enriched" assemblage characterized by high

numbers and biomasses of several deposit-feeding

species, especially the polychaetes Nephtys incisa and

Pherusa affinis and bivalve Nucula proxima. Tharyx spp.

and Cossura longocirrata were other numerically important polychaetes, and the phoronid worm Phoronis

architecta and anthozoan Ceriantheopsis americanus were

consistent dominants in both numbers and biomasses.

The assemblage was quite stable over the study period. There was an apparent tendency toward increasing abundance and dominance for Pherusa affinis.

More recent sampling for the sludge dumpsite "recovery" study indicates the continued persistence of the

assemblage, with P affinis remaining the biomass

dominant, at least into early 1989.

Components of the enriched assemblage were also

sometimes found in the sludge accumulation area,

and the assemblage was consistently present at all

other Christiaensen Basin stations, as well as at stations 2 and 3 immediately to the north of the Basin.

There were no clear changes over time at stations 2,

3, or 5, the CB station closest to the sludge area (2.8

km W of Station 6). There were, however, decreases

in overall faunal abundance at 9, 10, and 40.

Comparisons to Other Studies ------

The spatial pattern of assemblages in the Basin fits

the model of Pearson and Rosenberg (1978) for successional changes in response to organic inputs. Areas of high organic loading are often characterized by

large populations of a few small opportunistic species-this is represented by the Capitella spp.-dominated fauna at Station 6. Areas with somewhat lower

organic loading typically have high abundances of

several species, as in the enriched assemblage just described for most of the remaining Christiansen Basin

and some areas immediately outside the Basin.

Pearson and Rosenberg (1978) also note that sediments in the vicinity of a large organic input are

sometimes devoid of benthic macrofauna. No consistently defaunated zone has been found in the Basin;

for example, no azoic samples were collected in our

1979-85 surveys at Station 6, and only one of 40 grabs

in the intensive sampling of the sludge accumulation

area in the summers of 1986-1988 had no

macrofauna. The implication is that organic loading

rates in the Basin have not been as high, relative to

compensating phenomena such as dispersion and microbial breakdown, as in many other areas of organic

enrichment, e.g. the Saltkallefjord, Sweden; Loch

Creran, Scotland; Frazer River estuary, British Columbia; Kiel Bay, Germany (Pearson and Rosenberg

1978). This agrees with the Segar and Davis (1984)

global review of contaminated coastal areas, which

did not consider the New York Bight to be among the

most pollution-susceptible regions.

Detailed, quantitative comparisons of the 1979-87

data with results of earlier New York Bight surveys are

beyond the scope of this report. On a qualitative

level, there are clear long-term similarities. The en-
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 33

riched assemblage has been present in much of the

Basin at least since the first benthic surveys there in

1968 ( ational Marine Fisheries Service 1972; Pearce

1972; Pearce et al. 1981; Boesch 1982; Steimle et al.

1982; Steimle 1985). The Capitella spp.-dominated

fauna in the sludge accumulation area was not reported in the 1968-71 studies (National Marine Fisheries Service 1972; Pearce 1972). They were, however,

consistently found in the intensive 1973-75 MESA

sampling (Pearce et al. 1981). In that sampling, as in

ours, recurring high densities of Capitella spp. were

limited to a small area (the vicinity of Station 6). The

persistence of the enriched and Capitella spp. assemblages is evidence against gross changes in the

benthic environment of the Bight over the past 15-20

years. More sensitive measures such as numbers of

species and amphipods, however, do indicate that

there has recently been Widespread environmental

change in the Bight.

Summary

There were clear spatial patterns in the benthic

macrofauna of the New York Bight. Numbers of species and amphipods, considered relatively sensitive indicators of environmental quality, were lowest in the

"sludge accumulation area," followed by other

Christiaensen Basin and inshore Bight stations. Both

variables reached highest values at the outermost

shelf stations and in the outer Hudson Shelf Valley.

Cluster analysis of species composition also showed

spatial differences. This was expected, given the

ranges encountered in variables such as depth, substrate, and distance from estuaries and dumpsites.

There were several distinct assemblages, especially in

the sludge dumpsite, sludge accumulation area, midHudson Shelf Valley, and outer Shelf Valley. Long Island and New Jersey offshore stations were broadly

similar. All remaining Christiaensen Basin stations fell

into several related groups. The other inshore samplings were less similar to one another and were

spread over much of the dendrogram.

None of the variables examined distinguished

northern New Jersey inshore stations from stations

further south or along the Long Island coast. This

implies that the Hudson-Raritan plume, which tends

to follow the north Jersey coast and influence temperature, salinity, dissolved oxygen, carbon and toxicant levels there, has no overriding effect on the

benthos of that area.

The spatial patterns in numbers of species and amphipods were consistent between surveys. However,

absolute numbers of species and amphipods decreased at a large majority of stations between 1980

and 1985. Decreases for all stations combined were

highly significant (P= 0.0007 for species, 0.0004 for

amphipods). The most significant decreases in numbers of species and amphipods were in the Basin's

sludge accumulation area and at a site 52 km SE of

that area in the Hudson Shelf Valley. The latter site

had been chosen to represent the innermost part of

the Shelf Valley that had not been noticeably influenced by pollution. The pattern of decreases superficially suggests increasing effects of sludge or overall

waste loading, but the limited data available do not

indicate that waste inputs have increased, and there

are other feasible explanations for the observed

changes. At three inner Bight stations sampled more

recently, numbers of species generally rose somewhat

between 1986 and 1989, perhaps signalling a reversal

of the general downward trend from 1980 to 1985.

Temporal changes in species composition and dominants were less evident.

Spatial trends in numb_is of species and amphipods, and dominant species, were similar to those observed since sampling in the Christiaensen Basin area

began in 1968. A small area with a "highly altered"

assemblage dominated by the organic-enrichment indicator polychaetes Capitella spp. has been present in

the Basin since the early sampling, and an "enriched"

faunal assemblage with high biomasses of several species has consistently been found over most of the remaining basin. Samples devoid of benthic macrofauna have been very rare or absent in all surveys.

Gross changes in the benthic environment of the New

York Bight are therefore not evident over the past 15

years or more.

Acknowledgments-------- _

We thank Michelle Cox for the artwork. Dorothy

Jeffress and Joseph Vitaliano helped process the

1986-89 samples. Robert Pikanowski, Frank Steimle,

Joel O'Connor, Fred Grassle, Anne Studholme, and

two anonymous reviewers made helpful comments on

the manuscript.
34 NOAA Technical Report NMFS 103

Citations ---------------

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Green, R.H.

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Kuiper, j.

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Langton, R.W., and R.E. Bowman.

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Lee, W.Y., M.F. Welch, andJ.A.C. Nicol.

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29203,715 p.

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36:139-149.
36 NOAA Technical Report NMFS 103

LONG ISLAND OFFSHORE

33-

35-

,

I

I

-'

,

\

34-

" ,

,

I

,

38-

36·

37'

27-

26

"23

NEW

JERSEY

.----~7f~ n Dredged Material

LJ Dumpsite

IU?::q Sewag~ Sludge -:-:.:.>:.:. Dumpsite

Station locations in

Figure 1

ew York Bight, with subareas discussed in text. Diagonal lines indicate Christiaensen Basin.
Reid et at.: Benthic Macrofauna of the New York Bight, 1979-89 37

40Cl20'

NAUTICAL MILES

1 2 3 4 5

I I I I I I I I I I I I

.5

1

11111

I 2 4 6 8

KILOMETERS

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Basin

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Figure 2

Details of inner New York Bight.
38 NOAA Teehnieal Report NMFS 103

NEW

JERSEY

d ed Material OJ g~7n~site . . Sludge tI ~~~~~~e_~__

../

Figure 3 b r-Octobel- 1985. Numbers of specIe- sperO.l m.2 Septem e
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 39

I

o

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Numbers of species per 0.1 m2,July-August 1980.
...

0

Station 4

Number of grabs I z

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1001 I (')

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Figure 5

Means and 95% confidence limits for numbers of species per 0.1 m2 at each NEMP station on each sampling date, with numbers of replicate samples indicated.
Station

6

Number of grabs

5 5 5 5 5 5 5 52

55

5

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3 333

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90

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Figure5 (continued)

.,

cIQ' =F-t&)

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Number of grabs

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z

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Figure5 (continued)
N

Reid et aI.: Benthic Macrofauna of the New York Bight, 1979-89 43

CD

CD

(I)

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Station 26

Number of grabs

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79 80 81 82 83 84 85 86

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Figure5 (continued)
Station 31

Number of grabs

4

5 5 5 5

5

5

5 5 2

100

90

80

C'I 70- E,... 60 0

I a: I w

I Q. 50 I en

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Figure5 (continued)

:;I:l

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46 NOAA Technical Report NMFS 103

74°~ 1/ ,---------,

n Dredged Material

W Dumpsite

j:rr>1 ::::::::::::: Sewag~ DumpsiteSludge :p-""'----'

NEW

JERSEY

/

Figure 6

Numbers of amphipods per 0.1 m2, September-October 1985.
Reid et al.: Benthic Macrofauna of the New York Bight, 1979-89 47

NEW

JERSEY

o

o

Figure 7

Numbers of amphipods per 0.1 m2,July-August 1980.
""

00

Station 15 I

Number of grabs I z

0

5 5 5 5 5 5 5 5

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4000 -f t'l

[

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Figure 8

Means and 95% confidence limits for numbers of amphipods per 0.1 m2 at Station 15 in mid-Hudson Shelf Valley for each sampling date.

Numbers of replicate samples are indicated.
Figure 9

Dendrogram sho,\\,ing between-group similarities in species composition, with groups formed at 30% similarity levels. Each wider bar to left indicates the

level of similarity of samplings within a given group. Areas: NJI = New Jersey Inshore; NJO = New Jersl"Y Offshore; LII = Long Island Inshore; LIO = Long

Island Offshore; CB = Christiaensen Basin; HSV = Hudson Shelf Valley. n = number of samplings in group. Samplings in each group are given in Table 3.

Note log scale at negative similarities.

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Means and 95% confidence limits for numbers of Capitella sp. per 0.1 m2 at Station 6 ("sewage sludge accumulation area") for each sampling date. Numbers

of replicate samples are indicated.